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BULLETIN 
of the 


BRITISH 
ORNITHOLOGISTS’ CLUB 


EDITED BY 


Dr D. W. SNOW 
Soro CJ) PEARE 


Volume 117 
1997 


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ill 
LIST OF AUTHORS AND CONTENTS 
ALEIXO, A. Range extension of the Large-headed Flatbill Ramphotrigon 


megacephala with comments on its distribution ...................020-0-5 220 
ALSTROM. P., OLSSON, U. & COLSTON, P.R. Re-evaluation of the 
taxonomic status of Phylloscopus proregulus kansuensis Meise............... 177 


AMARILLA, L.A. see ERICSON, P.G.P.. 

AVERSA, T. see VALLELY, A.C. 

BARAHONA, O.R. see KRABBE, N. 

BARNES, R., BUTCHART, S.H.M., DAVIES, C.W.N., FERNANDEZ., M. 
& SEDDON, N. New distributional information on eight bird species from 
NINE TAR ios 5 5h RSE. /'etsid'k obeys AE shy to heat o-e)- Acce SP SOM 69 

BARNETT, J.M. see LOWEN, J.C. 

BARROWCLOUGH, G., LENTINO R., M. & SWEET, P.R. A new record of 


birds from Auyn-tepui, Estado Bolivar, Venezuela .....................-. 194 
Pape seen SCO! WTCED 325 es ks SK 2 es’ MS HE ONT Tce Ne 80, 151, 319 
BORGHESIO, L. Observations on the ecology of Tauraco ruspoli and T. leucotis 

MeEeneEn Pthionia.:.. .... 3%. £4 3eeCos--606 =H: R- DOA aa: Fh. 5! 11 


BOSISIO, A. see ORDANO, M. 
BRADSHAW, C.G., KIRWAN, G.M. & WILLIAMS, R.S.R. First record of 


Swainson’s Hawk Buteo swainsoni for the West Indies.................... 315 
BROOKS, T. & DUTSON, G. Twenty-nine new island records of birds from the 
INE SAM oe co en aes vs Ah che ciate OR RE SEE Sane. OT. 32 


BROOKS, T.M. see LOWEN, J.C. 
BUTCHART, S.H.M. see BARNES, R. 
CLANCEY, P.A. Variation in the Cape Penduline Tit Anthoscopus minutus of the 
i etETeMES: Fo 08 ks ee een ot LE PEM I. cag )-<b 52 
Subspeciation in Layard’s Tit-babbler of the southwestern Afrotropics ...... 244 
CLAY, R.P. see LOWEN, J.C. 
CLUB NOTICES 


Report of the Committee for 1996 & A.G.M. Agenda .................... 1 
Pee ESCM tL ACCTING. x. by on gpl. Tf As CESS LIL LO... Bf. 153 
Beametal statement for the year ended 31.12.96 . . 2.2. .angrbes cece reins - 161 
ic CHetwmrss <b atk A on: Mote ST LEI 58 ws BP Aa > oll 3, 81, 156, 241 


Joint Meeting between B.O.C. and the Linnean Society of London held on 
23.3.96 at Burlington House — see separate index at the end. 


VOLLARAN.J. he threat.status of the Sidamo'Larké 143.) sao. f.. Kh. CS OS 75 
COLSTON, P.R. see ALSTRM, P. 
Sere 4. Whaat is 2 desert bird? . ........ $4.2 TORT AD sae TRIAL 299 
DAVIES, C.W.N. see BARNES, R. 
DESFAYES, M. A specimen of Hirundo abyssinica from Madagascar .......... 315 


DICKERMAN, R.W. A substitute name for the Bioko race of Pycnonotus virens. v5 
DOMERGUE, C.A. see GOODMAN, S.M. 
DOWSETT, R.J. & MOORE, A. Swamp warblers Acrocephalus gracilirostris and 


anions a) | ake (Chad, Dliverdia 5 .9166140t- aeaa - baa: besedassdt do- ehees 48 
DUCKWORTH, J.W. Observations on a population of Jerdon’s Bushchat 
paascola jerdom in the Mekong channel, Laos ... ............i60. 5s «pase 210 


DUTSON , G. see BROOKS, T. 
ERICSON, P.G.P. Swedish records of the eastern Palearctic Hoopoe subspecies 


PP PHS SHINTBIA —- ss oe aes be Bs oe eA SAA OT eae. JAVA 19 
ERICSON, P.G.P. & AMARILLA, L.A. First observations and new 
@etbutioual data for birds in Paraguay ..... 42 - ASKER ane. 4M AS 60 


ESQUIVEL, E.Z. see LOWEN, J.C. see MADRONO N., A. 

FERNANDEZ, M. see BARNES, R. 

FLANAGAN, J.N.M. see TOYNE, E.P. 

FRITH, C.B. & FRITH, D.W. The taxonomic status of the bird of paradise 
Paradigalla carunculata intermedia (Paradisaeidae) with notes on the other 


EARPIECE ooo occ acess vaio sevintoviseamprezverarviese RBULIL Asoa.ne wlohe uhat 38 
FRITH, C.B. & FRITH, D.W. A distinctive new subspecies of Macgregor’s 
Bowerbird (Ptilonorhynchidae) of New Guinea................00.200005- 199 


FRITH, D.W. see FRITH, C.B. 
FROLANDER, A. see KRABBE, N. 


iv 


GALETTI, M., MARTUSCELLI, P., PIZO, M.A. & SIMAO, I. Records of 


Harpy and Creasted Eagles in the Brazilian Atlantic forest ................ 27 
GOMEZ DE SILVA G., H. Further observations on the nesting of the Azure- 
rumped: Tanager: 0.1)... 62. Se AL OO. 20 On Ee 16 
GONZALEZ M., O. First description of the nest of the Slender-billed Finch... 314 
GOODMAN, S.M., HAWKINS, A.F.A. & DOMERGUE, C.A. A new species 
of vanga (Vangidae, Calicalicus) from southwestern Madagascar ........... 5 
GREGORY, P. Range extensions and unusual sightings from Western Province, 
. Papua New Guinea. oi laa oo ae tu A ee 304 
GUSTEN, R. Fringilla coelebs gengleri — authorship and date revisited ........ 238 


HAWKINS, A.F.A. see GOODMAN, S.M. 
HAZEVOET, C.J. Ona record of the Wattled Crane Bugeranus carunculatus from 


Guinea-Bissau... .. 0.2: cc se acs oe ee ee 56 
HERZOG, S.K., KESSLER, -M., MAIJER, S.- & HOHNWALD, S: 
Distributional notes on birds of Andean dry valleys in Bolivia’)? !> S992) Be: 223 


HOHNWALD, S. see HERZOG, S.K. 

HOWELL, S.N.G. see LOUSADA, S.A. 

KESSLER, M. see HERZOG, S.K. 

KIRWAN, G.M. see BRADSHAW, C.G. 

KRABBE, N., POULSEN, B.O., FROLANDER, A. & BARAHONA. O.R. 
Range extensions of cloud forest birds from the high Andes of Ecuador: new 
sites for rare’or ‘little-recordéd' species 209 M200, FM St Se 248 

KRUL, R. see MORAES, V.S. 

LANUS, B.L.. see LOWEN, J.C. 

LENTINO R., M. see BARROWCLOUGH, G. 

see RODRIGUEZ, G. 

LILEY; D.C: see LOWEN, J.C. 

LOUSADA, S.A. & HOWELL, S.N.G. Amazona oratrix hondurensis: A new 
subspecies of parrot from the Sula Valley of northern Honduras .......... 203 

LOWEN, JC. (CEAY: Rees BARNEISE.. 0) vie MADRONO N, A., 
PEARMAN, M.., LANUS, B. L., TOBIAS, J.A., LILEY, EXC, BROOKS, 
Tavi. , ESQUIVEL, E.Z. & REID, J.M. New and noteworthy observations 
on Paraguayan avifauna® «oo... Se ©. 22 O08 TAU OE) 275 

MADRONO N., A. & ESQUIVEL, E.Z. Noteworthy records and range 
extensions from the Reserva Natural del Bosque Mbaracay (Mbaracay Forest 
Nature Reserve), Departamento de Canindey, Paraguay ..............-... 166 

MADRONO N., A. see LOWEN, J.C. 

MAIJER, S. see HERZOG, S.K. 

MARTUSCELLI, P. see GALETTI, M. 

MEES, G.F. On the identity of Heterornis senex Bonaparte ................... 67 

MOORE, A. see DOWSETT, R.J. 

MORAES, V.S. & KRUL, R. Notes on the Black-backed Tanager Tangara 
peruviana (Desmarest, 1806)........ 40.9 «SMES MA Se ee 316 

OLSSON, U. see ALSTROM, P. ORDANO, M. & BOSISIO, A. Historical 
records of threatened and near threatened Argentinian birds from Museo 
Provincial de Ciencias Naturales “‘Florentino Ameghino” of Santa Fe, 
Argentina»... ici nt teats BORE PSOE QUORS IN BIE Tt Se 77 

PARRINI, R. see RAPOSO, M.A. 

PATACHO, D. see REINO, L.M. 

PEARMAN, M. see LOWEN, J.C. 

PIZO, M.A. see GALETTI, M. 

POULSEN, B.O. see KRABBE, N. 


PRAWIRADILAGA, D.M. The Maleo Macrocephalon maleo on Buton ....... 237 
RAPOSO, M.A. & PARRINI, R. On the validity of the Half-collared Sparrow 
Arremon semitorquatus Swainson ........%.0. <b4. £02). 392 Be 4a eee 1837 294 


REID, J.M. see LOWEN, J.C. 
REINO, L.M. & PATACHO, D. A new beeding area for the Song Thrush 


Turdus philomelos in.north Portugal ...........0.1-c.eccn cine ee oe 312 
RODRIGUEZ, G. & LENTINO, M. Range expansion and summering of Palm 

Warbler Dendroica palmarum in Venezuela ..........500. 000 e0 02h ee 76 
SAGE, B. Melanism in the gulls (Laridae)..........%1.) 3244254. Soa 239 


SEDDON, N. see BARNES, R. 


SIMAO, I. see GALETTI, M. 

SMITH, P.W. The history and taxonomic status of the Hispaniolan Crossbill 
RES TENERIFE ss Soe ase ee lc AE EE RE ete oe eke alas wiles 

SWEET, P.R. see BARROWCLOUGH, G. 

TOBIAS, J.A. see LOWEN, J.C. 

TOYNE, E.P. & FLANAGAN, J.N.M. Observations on the breeding, diet and 
behaviour of the Red-faced Parrot Hapalopsittaca pyrrhops in southern 


VALLELY, A.C. & AVERSA, T. New and noteworthy bird records from Belize 

including the first record of Chestnut-collared Swift Cypseloides rutilus .... 
WALTERS, M. On the identity of Lophornis melaniae Floericke (Trochilidae) .. 
WILLIAMS, R.S.R. see BRADSHAW, C.G. 


AVIAN TAXONOMY FROM LINNAEUS TO DNA 
PAPERS PRESENTED AT A JOINT MEETING BETWEEN THE 
BRITISH ORNITHOLOGISTS’ CLUB AND THE LINNEAN 
SOCIETY OF LONDON HELD AT BURLINGTON HOUSE, 
23 MARCH 1996. 
BARBAGLI, Fabio, BARBAGLI, Fausto, & VIOLANI, C. Scopoli, Linnaeus 
aaa Wallcreeper Jichadroma muraria (V3 220098 CORSA AEE cam pyqeyl eae 

BARBAGLI, Fausto see BARBAGLI, Fabio 
see VIOLANI, C.G. 
Nh i eo .  e S ETP Ee oo oral mere et 
COLLAR, N.J. Taxonomy and conservation: chicken and egg ................ 
GALEB, B. see RUSSELL, D.A. 
GREENWOOD, J.J.D. Introduction: the diversity of taxonomies ............. 
HOATH, R. see RUSSELL, D.A. 
MIKHAILOV, K.E. Bird taxonomy based on eggshell structure .............. 
PARKIN, D. Molecular probes for identification of raptors.................6. 
RUSSELL, D.A., GALEB, B. & HOATH, R. X-raying the Gods: what were the 


SNOW, D.W. Should the biological be superseded by the phylogenetic species 
PAE ns a ilk a a SP AO © os meet cine 

VIOLANI, C.G. & BARBAGLI Fabio. Salvadori, Giglioli and Arrigoni: aspects 
of the trinomial controversy in Italian ornithology ................6..00.. 

VIOLANI, C.G. see BARBAGLI, Fabio. 

ee IES CONCCIIS 5.05 212 ecppeyaie ss ee ed ES, Mie a ed. 


264 


257 


272 
239 


145 
83 
122 
85 


149 
147 


148 
110 


137 


97 


INDEX TO SCIENTIFIC NAMES 
(Compiled by Mary N. Muller) 


All generic and specific names (of birds only) are indexed. New subspecific names are indexed 
in bold print under generic, specific and subspecific names. 


abyssinica, Coracias 82 

— , Hirundo 315 
Accipiter sp. 32 

— bicolor 233, 272 

— erythronemius 168 

— poliocephalus 306 

— poliogaster 279 

— superciliosus 279 
accipitrinus, Deroptyus 126 
Aceros leucocephalus 127-8 

— waldeni 127-8 
Acestrura mulsant 234 
Acrocephalus arundinaceus 310 

— brevipennis 118, 156—7 

— gracilirostris 48—50 

— rufescens 48-50 
Actitis hypoleucos 33 

— miacularia 196, 198, 233 
acuticaudata, Aratinga 232, 233 
adela, Oreotrochilus 234 
adusta, Premnoplex 198 
aedon, Troglodytes 234 
Aegolius harrisii 196, 198, 228, 234 
aenea, Chloroceryle 172, 284 
aequinoctialis, Geothlypis 235 
Aeronautes andecolus 234 

— montivagus 198,234 
aeruginosus, Circus 149 
aestiva, Amazona 234 
aethereus, Nyctibius 171 

— , Phaethon 157 
affinis, Apus 35 

— , Phylloscopus 186 
africanus, Buphagus 4 
Agelaius flavus 65 
Alaemon sp. 156 
Alauda razae 118, 156-7 
alba, Ardea 233 

— , Strix 137 

— , Tyto 156, 242 
albicaudatus, Buteo 233 
albicollis, Nyctidromus 234 
albifrons, Branta 137 
albigula, Buteo 233 
albigularis, Rhinomyias 128 
albinucha, Xenopsaris 65, 231, 234 
albitarsus, Ciccaba 251, 253 
albocristata, Sericossypha 255 
albofrontatus, Heterornis 67 

— , Sturnus 68 

— , Temenuchus 67 
albogularis, Otus 253 
albolimbatus, Megalurus 310 
albolineatus, Lepidocolaptes 234 
albonotataus, Buteo 233 
Alectoris philbyi 82 


Alectrurus risora 64, 78 
alexandri, Apus 118 
aluco, Strix 34 
amadoni, Andropadus virens nom.nov. 
75 
amaurocephalus,. Leptopogon 234 
amaurochalinus, Turdus 232 235 
Amaurospiza concolor 273 
amaurotis, Anabacerthia 284-5, 290 
Amazilia chionogaster 232, 234 
— tobaci 197-8 
— versicolor 197-8 
Amazona aestiva 234 
— auropalliata 209 
— ochrocephala 205-9 
— oratrix 205-9 
Amazona oratrix hondurensis subsp. 
nov. 205-9 
— pretrei 282, 290 
— vinacea 282 
amazona, Chloroceryle 234 
amazonina, Hapalopsittaca 257 
Amblyornis inornatus 46 
— macgregoriae 199-204 
Amblyornis macgregoriae lecroyae 
subsp.nov. 201-4 
americana, Anas 272 
— Chloroceryle 234 
— , Rhea 78 
amethysticollis, Heliangelus 249 
amethystina, Calliphlox 228, 234 
Ammodramus aurifrons 235 
Ammomanes sp. 156 
— cincturus 301 
— deserti 301 
amurensis, Falco 4 
Anabacerthia amaurotis 284-5, 290 
Anairetes flavirostris 234 
analis, Catamenia 235 
Anas americana 272 
— aucklandica 127 
— cyanoptera 272 
— discors 272 
— querquedula 309 
andecolus, Aeronautes 234 
andicola, Leptasthenura 252 
Andropadus gracilirostris 75 
Andropadus virens amadoni nom.nov. 
15 
angustifrons, Psarocolius 235 
angustirostris, Lepidocolaptes 232, 234 
ani, Crotophaga 232, 234 
Anodorhynchus hyacinthinus 160, 170 
anomalus, Eleothreptus 283 
Anseranas semipalmatus 309 
antarcticus, Anthus 160 Cinclodes 242 


Anthoscopus caroli 52-5, 157-8 

— minutus 52-5 
Anthus antarcticus 160 

— hellmayri 275, 278, 289 

— nattereri 286, 289, 290 

— rubescens 120 

— rufulus 213 
Anurolimnas viridis 275, 276, 289 
apiaster, Merops 82 
Aplonis metallica 308 

— mystacea 308 
apoda, Paradisaea 141, 309 
Apus affinis 35 

— alexandri 118 

— pacificus 35 
Aquila pomarina 149 
aquila, Eutoxeres 69 
Ara auricollis 233 

— chloroptera 196, 198 

— militaris 233 

— rubrogenys 227, 233 

— severa 233 
Aramides cajanea 233 

— wolfi 160 
Aratinga acuticaudata 232, 233 

— leucophthalmus 69 

— mitrata 233 
arcuata, Dendrocygna 32 

— , Pipreola 252 
Ardea alba 233 

— cocoi 233 

— goliath 82 

— purpurea 156 
ardeola, Dromas 82 
Ardeotis kori 4 
arfakiana, Melanocharis 305-6 
argentatus, Larus 239 
armandii, Phylloscopus 186 
armata, Merganetta 233 
armillata, Cyanolyca 250 
aroyae, Thamnophilus 234 
Arremon flavirostris 235, 295-6 

— semitorquatus 294-8 

— taciturnus 294-8 
arundinaceus, Acrocephalus 310 
assimilis, Puffinus 118, 157 
Asthenes berlepschi 229, 234 

— dorbignyi 230, 234 
Astrapia nigra 46 
ater, Parus 114—-5 
aterrimus, Knipolegus 234 
Atlaptes fulviceps 235 

— personatus 198 

— torquatus 235, 
atra, Tijuca 243 
atratus, Coragyps 195, 198, 233 
atricapilla bigilalei, Lonchura 4 

— moresbyi, Lonchura 4 

— obscura, Lonchura 4 

— selimbauensis, Lonchura 4 
atricapillus, Herpsilochmus 234 
atricilla, Larus 240 


atrocapillus, Crypturellus 233 
atrovirens, Psarocolius 235 
Atticora fasciata 234 
aucklandica, Anas 127 
Aulacorhynchus prasinus 250 
aura, Cathartes 195, 198, 233 
aurantiirostris, Saltator 65, 235 
aurantius, Chaetops 245 
aureoventris, Chlorostilbon 232, 234 

— , Pheucticus 232, 235 
aureus, Sericulus 309 
auricollis, Ara 233 
auriculata, Zenaida 233 
aurifrons, Ammodramus 235 

— , Neopelma 125 
auropalliata, Amazona 209 
australe, Dicaeum 128 
australis, Coturnix 304 

— Phalcoboenus 242 
Automolus roraimae 198 
aymara, Bolborhynchus 233 
azarae, Synallaxis 234 
azurea, Hypothymis 36 


badius, Molothrus 235 
bannermani, Buteo 156 
baroni, Metallura 254 
Basileuterus bivittatus 235 

— coronatus 235 
Batara cinerea 234 
beccarii, Cochoa 125 
berlepschi, Asthenes 229, 234 

— , Thripophaga 71 
biarmicus, Falco 149 
bicolor, Accipiter 233, 272 
bifasciata, Loxia 264-9 
bigilalei, Lonchura atricapilla 4 
bivittatus, Basileuterus 235 
blakei, Grallaria 72 
blanfordi, Pycnonotus 213 
boehmi, Parisoma 244 
boissonneautii, Pseudocolaptes 258 
Bolborhynchus aymara 233 
boliviana, Poospiza 235 
bolivianus, Oreopsar 235 

— , Zimmerius 234 
bonapartei, Nothocercus 248 
bonariensis, Thraupis 235 
bradfieldi, Tockus 4 
brachydactyla, Tachyeres 242 
brachyurus, Buteo 233 
branickii, Odontorchilus 71 
Branta albifrons 137 
brasiliana, Hydropsalis 234 
brasilianum, Glaucidium 234 
brasilianus, Phalacrocorax 233 
brevicauda, Paradigalla 38—47 


brevipennis, Acrocephalus 118, 156-7 


breweri, Spizella 130 

brissonii, Cyanocompsa 65, 235 
Brotogeris versicolurus 227, 233 
bruyjnii, Micropsitta 306 


brunneinucha, Buarremon 251 
brunniceps, Myioborus 232, 235 
Buarremon brunneinucha 251 
torquatus 251 

Bubo philippensis 34 
virginianus 234 

Bucanetes githagineus 82, 301 
Bucorvus leadbeateri 4 
buccinator, Cygnus 129 
budytoides, Stigmatura 232, 234 
Bugeranus carunculatus 4, 56—9 
Buphagus africanus 4 
burmeisteri, Phyllomyias 234 
buryi, Parisoma 244 

Buteo albicaudatus 233 

albigula 233 

albonotataus 233 
bannermani 156 
brachyurus 233 
jamaicensis 316 
magnirostris 195, 198, 232, 233, 249 
polyosoma 233, 258 
swainsoni 281, 290, 315-6 
Buteogallus urubitinga 195, 198 
Buthraupis eximia 73 

wetmorei 255 

Butorides striatus 82 


cabanisi, Tangara 16-8 

Cacatua galerita 126 

sulphurea 126 

Cacicus chrysopterus 235 
leucorhamphus 258 
Cacomantis merulinus 213 
caerulescens, Sporophila 235 

, Chamnophilus 234 
caeruleus, Cyanerpes 235 

, Elanus 149 

cafer, Promerops 245 

cajanea, Aramides 233 

Calicalicus madagascariensis 5—10 
Calicalicus rufocarpalis sp.nov. 5-10 
Calliphlox amethystina 228, 234 
Calonectris spp. 118 

diomedea 157 

Calyptura cristata 243 

camelus, Struthio 4 

campanisona, Chamaeza 234 
campbelli, Malurus 204, 308 
Campephilus leucopogon 234 
melanoleucos 234 
rubricollis 234 
Camptostoma obsoletum 234 
Campylopterus hyperythrus 198 
largipennis 234 
Campylorhamphus falcularius 285, 290 
pucheranu 255 
trochilirostris 234 
canadensis, Sitta 140 

canaria, Serinus 180 

candicans, Caprimulgus 275, 277, 289, 290 
canus, Picus 127 


Vill 


capensis, Zonotrichia 198, 235, 258 
capnodes, Otus 125 

caprata, Saxicola 213, 304 
Caprimulgus candicans 275, 277, 289, 290 
longirostris 198, 228, 234 
maculicaudus 275, 277, 289, 290 
rufus 234 

sericocaudatus 171 

carbo, Ramphocelus 173, 235 
Carduelis sp. 235 

magellanica 235 
xanthogastra 235 

Cariama cristata 170 

caripennis, Steatornis 196, 198 
caroli, Anthoscopus 52-5, 157-8 
Carpococcyx radiatus 126 
viridis 126 

Carpornis cucullatus 243 
carunculata, Paradigalla 38-47 
carunculatus, Bugeranus 4, 56-9 
Casiornis rufas 234 
castaneocapillus, Myioborus 198 
castaneus, Pachyramphus 234 
castanotis, Pteroglossus 234 
Catamenia analis 235 
homochroa 198 

Cathartes aura 195, 198, 233 
caudacuta, Culicivora 172 
caudatus, Theristicus 232-3 
cayana, Piaya 234 

, Llityra 278 

cayanensis, Icterus 231, 235 
cayennensis, Panyptila 234 
celebensis, Hirundapus 35 
Centropus sinensis 213 
cephalotes, Myiarchus 234 
Certhilauda chuana 4 

cerulea, Dendroica 273 

Ceryle torquata 234 

chacuru, Nystalus 234 

Chaetops aurantius 245 

frenatus 245 

Chaetura cinereiventris 171 
vauxi 272 

chalybea, Euphonia 287, 290 
Chamaeza campanisona 234 
mollissima 71-2, 250 
Charadrius collaris 232, 233 
chiguanco, Turdus 235 
chinchipensis, Synallaxis 125 
chinensis, Streptopelia 34 
chionogaster, Amazilia 232, 234 
chiriquensis, Elaenia 197-8, 280, 289 
Chloephaga hybrida 242 

picta 241 

rubidiceps 241 
Chloroceryle aenea 172, 284 
amazona 234 

americana 234 

inda 171 

chloropetoides, Thamnornis 8 
Chloropipo uniformis 198 


chloroptera, Ara 196, 198 
Chlorornis riefferii 251 
Chlorospingus ophthalmicus 235 
Chlorostilbon aureoventris 232, 234 

— mellisugus 234 
chlorotica, Euphonia 235 
choliba, Otus 228, 234 
Chondrohierax uncinatus 62, 168, 233 
chrysocephalus, Myiodynastes 234 
chrysochloros, Piculus 234 
Chrysocolaptes lucidus 127 
chrysops, Cyanocorax 235 

— , Zimmerius 73 
chrysopterus, Cacicus 235 
chuana, Certhilauda 4 
Ciccaba albitarsus 251, 253 

— huhula 62-3 

— nigrolineata 63 

— virgata 171 
Ciconia ciconia 4 
cinchoneti, Conopias 224, 230, 234 
Cinclodes antarcticus 242 

— fuscus 234 
Cinclus leucocephalus 234 
cincturus, Ammomanes 301 
cinerea, Batara 234 

— , Serpophaga 234 
cinereicapillus, Zimmerius 73 
cinereiventris, Chaetura 171 
cinereum, Conirostrum 231, 235 

— , Todirostrum 234 
cinereus, Circus 281 

— , Contopus 234 
cinnamomea, Pyrrhomyias 234, 258 

— , Sporophila 174, 288, 290 


cinnomomeiventris, Ochthoeca 250, 252 


Cinnycerthia unirufa 252 
Circus spp. 149 

— aeruginosus 149 

— cinereus 281 

— macrourus 4 

— pygargus 4 
cirratus, Picumnus 234 
Cissopis leveriana 235 
Cisticola exilis 213 
Claravis godefrida 174, 279, 290 

— mondetoura 227, 233 

— pretiosa 279 


Clibanornis dendrocolaptoides 284, 290 


Cnemotriccus fuscatus 234 
cobbi, Troglodytes 242 
Coccothraustes coccothraustes 140 
Cochoa beccarii 125 
cocoi, Ardea 233 
coelebs, Fringilla 238-9 
coelestis, Hypothymis 36 
Coeligena lutetiae 252 
Coereba flaveola 173, 198, 253 
Colaptes melanochloros 228, 234 
Colibri coruscans 198 

— delphinae 196, 198 

— serrirostris 234 


— thalassinus 234 
collaris, Charadrius 232, 233 

— , Trogon 234 
Collocalia sp. 34-5 

— esculenta 35 

— fuciphaga 35 

— germani 35 

— mearnsi 34-5 

— papuensis 304 

— vanikorensis 35 
Colluricincla harmonica 305 

— megarhynchus 308 
collurio, Lanius 4 
collybita, Phylloscopus 160 
Colonia colonus 234 
colonus, Colonia 234 
Columba fasciata 198, 227, 233 

— miaculosa 233, 282, 288 

— plumbea 233 

— speciosa 170 
Columbina picui 233 

— talpacoti 233 
comrii, Manucodia 129 
concolor, Amaurospiza 273 

— , Xenospingus 314 
Conirostrum cinereum 231, 235 

— speciosum 231, 235 
Conopias cinchoneti 224, 230, 234 
Contopus cinereus 234 

— fumigatus 234, 250 
coprotheres, Gyps 4 
Copsychus saularis 213 
Coracias abyssinica 82 
Coracopsis nigra 159 

— vasa 159 
Coragyps atratus 195, 198, 233 
coronatus, Basileuterus 235 

— , Harpychaliaetus 77 

— , Pterocles 301 
coruscans, Colibri 198 
Corvus ruficollis 301 
Coryphaspiza melanotis 79, 281, 290 
Coryphospingus cucullatus 235 
Coturnicops notata 78 
Coturnix australis 304 
Cranioleuca sp. 229, 234 

— demissa 198 

— pyrrhophia 234 
cristata, Calyptura 243 

— , Cariama 170 
cristatellus, Cyanocorax 173, 289 
Crotophaga ani 232, 234 
Crypturellus atrocapillus 233 

— obsoletus 233 

— ptaritepui 195, 198 

— tataupa 233 

— undulatus 233 
cucullatus, Carpornis 243 

— , Coryphospingus 235 
Culicivora caudacuta 172 
cumingii, Megapodius 32 
cursor, Cursorius 156, 301 


Cursorius cursor 156, 301 
curucui, Trogon 234 
curvirostra, Loxia 264-9 
Cyanerpes caeruleus 235 
cyanocephala, Euphonia 235 
Cyanocompsa brissonii 65, 235 
Cyanocorax chrysops 235 

— cristatellus 173, 289 

— cyanomelas 235 
Cyanolanius madagascarinus 5 
cyanoleuca, Notiochelidon 234 
Cyanolyca armillata 250 

— turcosa 250-1, 258 
cyanomelas, Cyanocorax 235 
cyanoptera, Anas 272 

— , Tangara 198 
Cyclarhis gujanensis 235, 251 
Cygnus buccinator 129 
Cypseloides phelpsi 196, 198 

— rutilus 234, 272-3 


dactylatra, Sula 81 
davisoni, Phylloscopus 186 
dayi, Elaenia 198 
decumanus, Psarocolius 235 
delattrei, Lophornis 235-6 
delphinae, Colibri 196, 198 
demissa, Cranioleuca 198 
Dendrocopos dorae 82 
Dendroica cerulea 273 

— palmarum 76 

— striata 77 
Dendrocolaptes picumnus 234 


dendrocolaptoides, Clibanornis 284, 290 


Dendrocygna sp. 32 

— arcuata 32 

— guttata 32 
derbianus, Oreophasis 16 
Deroptyus accipitrinus 126 
deserti, Ammomanes 301 

— , Oenanthe 301 
desmarestii, Psittaculirostris 307 
diadematus, Stephanophorus 287, 290 
diamondi, Phonygammus 130-1 
Dicaeum australe 128 

— haematostictum 125, 128 

— pectorale 305 
Dicrurus hottentottus 127 
Diglossa major 198 

— sittoides 235 
Diomedea exulans 160 
diomedea, Calonectris 157 
discors, Anas 272 
doliatus, Thamnophilus 234 
dominicus, Podiceps 198 
dorae, Dendrocopos 82 
dorbignyi, Asthenes 230, 234 
dorsalis, Mimus 235 
Dromas ardeola 82 
Dromococcyx phasianellus 170 
Drymophila rubricollis 285, 290 
Dryocopus galeatus 78 


— lineatus 234 

— schulzi 79 
dumicola, Polioptila 232, 234 
duvaucelii, Vanellus 213 


ecaudatus, Terathopius 4, 82 
Eegretta gularis 82 

— thula 233 

— vinaceigula 4 
Elaenia chiriquensis 197-8, 280, 289 

— dayi 198 

— obscura 234 

— pallatangae 198 
Elanoides forficatus 168, 195, 198, 233 
Elanus caeruleus 149 
Electron platyrhynchum 234 
elegans, Laniisoma 243 
elegantissima, Euphonia 18 
Eleothreptus anomalus 283 
eliciae, Hylocharis 273 
Emberizoides ypiranganus 289, 290 
Embernagra platensis 232, 235 
emeiensis, Phylloscopus 180 
Empidonax sp. 234 
Entomodestes leucotis 235 
Ephippiorhynchus senegalensis 4, 57 
epops, Upupa 19-26, 82 
Eremopterix sp. 156 

— nigriceps 301 
erythronemius, Accipiter 168 
esculenta, Collocalia 35 
Eudynamys scolopacea 34 
euleri, Lathrotriccus 234 
Euphonia sp. 235 

chalybea 287, 290 

~ chlorotica 235 

— cyanocephala 235 

— elegantissima 18 

— laniirostris 235 
Eurylaimus samarensis 126 

— steeru 126 
euryptera, Opisthoprora 254 
Eutoxeres aquila 69 
exilis, Cisticola 213 

— , Ixobrychus 281, 288 

— , Laterallus 282 
eximia, Buthraupis 73 
exulans, Diomedea 160 


falcinellus, Plegadis 309 
falcirostris, Sporophila 173, 287 
Falco amurensis 4 

— biarmicus 149 

— fermoralis 233 

— naumanni 4 

— pelegrinoides 149 

— peregrinus 32, 156, 233 

— rufigularis 233 

— sparverius 233 

— tinnunculus 149, 156 

— , vespertinus 4 
falcularius, Campylorhamphus 285, 290 


fasciata, Atticora 234 

— , Columba 198, 227, 233 
fasciatum, Tigrisoma 233 
fasciatus, Myiophobus 64, 234 
fasciicauda, Milvus 156 
feae, Pterodroma 129, 157 
femoralis, Falco 233 
ferox, Myiarchus 234 
ferruginea, Hirundinea 198, 234 
finlaysoni, Pycnonotus 213 
flava, Motacilla 310 

— , Piranga 235 
flaveola, Coereba 173, 198, 253 

— , Sicalis 235 
flavigaster, Microeca 305 
flavipes, Notiochelidon 254 
flavirostris, Anairetes 234 

— , Arremon 235, 295-6 

— ,Rynchops 4 
flaviventris, Prinia 212 
flavus, Agelaius 65 
fluviatilis, Locustella 4 
forficatus, Elanoides 168, 195, 198, 233 
Formicivora melanogaster 234 

— rufa 63 
Fregata magnificens 157 
frenatus, Chaetops 245 
Fringilla coelebs 238-9 
frontalis, Synallaxis 234 

— , Veniliornis 234 
fuciphaga, Collocalia 35 
fulgidus, Psittrichas 307 
fulica, Heliornis 169 
fuliginosa, Tiaris 175, 288 
fuliginosus, Haematopus 158, 243 
fulviceps, Atlapetes 235 
fumigatus, Contopus 234, 250 

— , Veniliornis 234 
furcata, Thalurania 234 
Furnarius rufus 232, 234 
fusca, Porzana 33 
fuscata, Sterna 81 
fuscater, Turdus 235, 258 
fuscatus, Cnemotriccus 234 
fuscus, Cinclodes 234 

— , Larus 239 

— , Melanotrochilus 283 


gabar, Micronisus 307 
gaimardii, Myiopagis 234 
galatea, Tanysiptera 307 
galbula, Ploceus 82 
galeatus, Dryocopus 78 
galertia, Cacatua 126 
Gallinago imperialis 252 

— paraguaiae 198 
Gallinula melanops 288, 290 
Garrulus glandarius 140 
genibarbis, Thryothorus 234 
Geopelia striata 34, 304 
Geothlypis aequinoctialis 235 
Geranoaetus melanoleucus 232, 233 


Xi 


germani, Collocalia 35 
gigantea, Melampitta 305 
gigas, Patagona 234 
githagineus, Bucanetes 82, 301 
glandarius, Garrulus 140 
Glareola lactea 213 

— maldivarum 33 

— nordmanni 4 
glareola, Tringa 33 
Glaucidium sp. 234 

— brasilianum 234 
godefrida, Claravis 174, 279, 290 
goiavier, Pycnonotus 36 
goliath, Ardea 82 
Gorsachius leuconotus 4 
Goura scheepmakeri 307 
gracilirostris, Acrocephalus 48—50 

— , Andropadus 75 
Grallaria blakei 72 

— przewalskii 72 
Grallaricula lineifrons 254 

— nana 254 
grayi, Locustella 305 

— , Malurus 204, 308 
griseicapillus, Sittasomus 234 
griseisticta, Muscicapa 305 
griseocristatus, Lophospingus 232, 235 
griseolus, Phylloscopus 186 
griseus, Nyctibius 234 
Grus grus 82 
gryphus, Vultur 233 
guianensis, Morphnus 27-30 
guira, Hemithraupis 235 
gujanensis, Cyclarhis 235, 251 
gularis, Egretta 82 

— , Rhinomyias 128 
gurneyi, Promerops 245 
guttata, Dendrocygna 32 

— , Ortalis 233 
guttatus, Xiphorhynchus 234 
Gyps coprotheres 4 


haemastica, Limosa 78 
haematodus, Trichoglossus 127 
Haematopus fuliginosus 158, 243 
haematostictum, Dicaeum 125, 128 
Halcyon leucocephala 118 
Haliaeetus leucogaster 32 
Hapalopsittaca amazonina 257 

— melanotis 262 

— pyrrhops 253, 257-62 
harmonica, Colluricincla 305 
Harpia harpyja 27-30, 78 
Harpyhaliaetus coronatus 77 

— solitarius 233 
harpyja, Harpia 27-30, 78 
harrisii, Aegolius 196, 198, 228, 234 
harterti, Upucerthia 229, 234 
helenae, Hypothymis 36 
Heliangelus amethysticollis 249 

— regalis 69 
Heliodoxa xanthogonys 196, 198 


Heliomaster longirostris 234 
Heliornis fulica 169 
hellmayri, Anthus 275, 278, 289 
Hemiprocne longipennis 34 
Hemithraupis guira 235 
Hemitriccus margaritaceiventer 234 
Herpsilochmus atricapillus 234 
Heteromirafra sidamoensis 15, 75 
Heterornis albofrontata 67 

— senex 67-8 
Hieraaetus pennatus 316 
Hippolais icterina 4 

— olivetorum 4 
Hirundapus celebensis 35 
Hirundinea ferruginea 198, 234 
Hirundo abyssinica 315 

— megaensis 15 

— rustica 36 

— smithii 213 
hirundo, Sterna 33 
hispanica, Oenanthe 142 
hodgsonii, Prinia 213 
homochroa, Catamenia 198 


hondurensis, Amazona oratrix subsp. 


nov. 205-9 

hornbyi, Oceanodroma 159 
hottentottus, Dicrurus 127 
hoyi, Otus 228, 234 
huhula, Ciccaba 62-3 

— , Strix 283 
hyacinthinus, Anodorhynchus 160, 170 
hybrida, Chloephaga 242 
hydrocharis, Tanysiptera 307 
Hydropsalis brasiliana 234 
Hylocharis eliciae 273 

— sapphirina 283-4 
Hylopezus nattereri 280, 290 
hylophila, Strix 283 
Hylophilus hypoxanthus 235 
hyperythrus, Campylopterus 198 
hypochroma, Sporophila 174 
hypoleucos, Actitis 33 
hypostictus, Taphrospilus 234 
Hypothymis azurea 36 

— coelestis 36 

— helenae 36 
hypoxanthus, Hylophilus 235 


iagoensis, Passer 118, 156 
icterina, Hippolais 4 
icterophrys, Satrapa 234 
Icterus cayanensis 231, 235 
— icterus 66 
Ictinia plumbea 233 
ignicapillus, Regulus, 160 
ignobilis, Turdus 197-8 
iliaca, Passerella 101-5 
iliacus, Turdus 140 
impennis, Pinguinus 159 
imperialis, Gallinago 252 
imthurni, Macroagelaius 198 
incerta, Pitohui 308 


inda, Chloroceryle 171 
inepta, Megacrex 307 
inornata, Prinia 213 
inornatus, Amblyornis 46 

— , Rhabdornis 36 
inquieta, Myiagra 310 
inquisitor, Tityra 278 
insignis, Thamnophilus 198 
intermedia, Paradigalla 39-47 
isabellina, Oenanthe 82 
isabellinus, Lanius 82 
Ixobrychus exilis 281, 288 

— minutus 309 


Jabiru mycteria 160 
jacarina, Volatinia 235 
jacquacu, Penelope 233 
jacutinga, Pipile 78 
jamaicensis, Buteo 316 
jerdoni, Saxicola 210-8 


keiensis, Micropsitta 306 
Knipolegus aterrimus 234 

— poecilurus 197-8 
koeniswaldiana, Pulsatrix 282 
kori, Ardeotis 4 


lactea, Glareola 213 
laniirostris, Euphonia 235 
Laniisoma elegans 243 
Lanius collurio 4 

— isabellinus 82 

— minor 4 
largipennis, Campylopterus 234 
Larus argentatus 239 

— atricilla 240 

— fuscus 239 

— leucophthalmus 82 

— ridibundus 239 
lateralis, Poospiza 281, 290 
Laterallus exilis 282 
Lathrotriccus euleri 234 
layardi, Parisoma 244-7 
leachii, Mackenziaena 280, 290 
leadbeateri, Bucorvus 4 
lecroyae, Amblyornis macgregoriae 

subsp.nov. 201-4 

Legatus leucophaius 234 
Lepidocolaptes albolineatus 234 

— angustirostris 232, 234 
Leptasthenura andicola 252 
Leptopogon amaurocephalus 234 

— superciliaris 234 
Leptotila megalura 232, 233 

— verreauxi 233 
leucocephala, Halcyon 118 
leucocephalus, Aceros 127-8 

— , Cinclus 234 
leucogaster, Haliaeetus 32 

— , Sula 81, 157 
leuconota, Pyriglena 234 
leuconotus, Gorsachius 4 


leucophaius, Legatus 234 
leucophrys, Mecocerculus 198, 234 
, Ochthoeca 234 

, Vireo 251 
leucophthalmus, Aratinga 69 

, Larus 82 

leucopogon, Campephilus 234 
leucoptera, Loxia 264~-9 
leucorhamphus, Cacicus 258 
leucoryphus, Platyrinchus 64 
leucosticta, Lonchura 308 
leucotis, Entomodestes 235 

, Tauraco 11-5 
leveriana, Cissopis 235 
lherminieri, Puffinus 118 
lichtensteinii, Pterocles 301 
lignarius, Picoides 228, 234 
Limosa haemastica 78 
lineatus, Dryocopus 234 
lineifrons, Grallaricula 254 
Lochmias nematura 198 
Locustella fluviatilis 4 

grayi 305 

Lonchura atricapilla bigilalei 4 
atricapilla moresbyi 4 
atricapilla obscura 4 
atricapilla selimbauensis 4 
leucosticta 308 
nevermanni 310 
punctulata 36 

striata sumatrensis 4 
stygia 310 

vana 46 

longicauda, Melanocharis 307 
, Myrmotherula 234 
longipennis, Hemiprocne 34 
longirostris, Caprimulgus 198, 228, 234 
, Heliomaster 234 
Lophornis delattrei 235-6 
melaniae 235-6 
stictolopha 235-6 
Lophospingus griseocristatus 232, 235 
Loxia bifasciata 264-9 
curvirostra 265-9 
leucoptera 264~-9 
megaplaga 264-70 
lubomirsku, Pipreola 72-3 
lucidus, Chrysocolaptes 127 
lugens, Parisoma 244 
Lurocalis rufiventris 251, 253 
luteola, Sicalis 274 

lutetiae, Coeligena 252 
Lycocorax pyrrhopterus 119 
lyra, Uropsalis 234 


macgregoriae, Amblyornis 199-204 

macgregoriae lecroyae, Amblyornis 
subsp.nov. 201-4 

Machetornis rixosus 234 

Mackenziaena leachii 280, 290 

Macroagelaius imthurni 198 

Macrocephalon maleo 237 


Xill 


macrourus, Circus 4 

macularia, Actitis 196, 198, 233 

maculatus, Myiodynastes 234 

, Nystalus 234 

maculiacaudus, Caprimulgus 
289, 290 

maculirostris, Muscisaxicola 230, 234 

maculosa, Columba 233, 282, 288 

madagascariensis, Calicalicus 5-10 

madagascarinus, Cyanolanius 5 

magellanica, Carduelis 235 

magnificens, Fregata 157 

magnirostris, Buteo 195, 198, 232, 233, 
249 

major, Diglossa 198 

, Pachyramphus 273 

, Taraba 234 

, Tinamus 233 

, Xiphocolaptes 234 

maldivarum, Glareola 33 

maleo, Macrocephalon 237 

Malurus campbelli 204, 308 

gray1 204, 308 

Manucodia comrii 129 

margaritaceiventer, Hemitriccus 234 

markhami, Oceanodroma 159 

maximiliani, Melanopareia 232, 234 

, Pionus 234 

maximus, Saltator 235 

mearnsi, Collocalia 34-5 

Mearnsia picina 35 

Mecocerculus leucophrys 198, 234 

megacephala, Ramphotrigon 172, 220-3 

Megacrex inepta 307 

megaensis, Hirundo 15 

megalopterus, Phalcoboenus 233 

megalura, Leptotila 232, 233 

Megalurus albolimbatus 310 

timoriensis 304, 310 

megaplaga, Loxia 264-70 

Megapodius cumingii 32 

megarhynchus, Colluricincla 308 

, Rhamphomantis 307 

Melampitta gigantea 305 

melancholicus, Tyrannus 234 

melaniae, Lophornis 235-6 

Melanocharis sp. 305-6 

arfakiana 305-6 

longicauda 306 

striativentris 306 

melanochloros, Colaptes 228, 234 

Melanodera melanodera 242 

melanogaster, Formicivora 234 

melanoleuca, Poospiza 235 

melanoleucos, Campephilus 234 

melanoleucus, Geranoaetus 232, 233 

, Spizastur 168 

melanonota, Pipraeidea 231, 235 

Melanopareia maximiliani 232, 234 

torquata 172, 286, 289 

melanopis, Schistochlamys 235 

melanops, Gallinula 288, 290 


Diana slain 


melanotis, Coryphaspiza 79, 281, 290 

— , Hapalopsittaca 262 
Melanotrochilus fuscus 283 
mellisugus, Chlorostilbon 234 
menstruus, Pionus 234 
Merganetta armata 233 
Merops apiaster 82 

— philippinus 213 
merula, Turdus 112-4 
merulinus, Cacomantis 213 
metallica, Aplonis 308 
Metallura baroni 254 

— odomae 254 

— williami 254 
Microeca flavigaster 305 
Micronisus gabar 307 
Micropsitta bruijnii 306 

— keiensis 306 


Micropygia schomburgkii 275, 277, 289 


militaris, Ara 233 
milleri, Polytmus 198 
Miulvus fasciicauda 156 

— milvus 83, 147-8 
Miumus dorsalis 235 
minor, Lanius 4 

— , Paradisaea 119 

— , Phoenicopterus 4 
minutus, Anthoscopus 52-5 

— , Ixobrychus 309 
mitrata, Aratinga 233 
Mniotilta varia 77 
modestus, Sublegatus 232, 234 
molinae, Pyrrhura 233 
mollissima, Chamaeza 71-2, 250 
Molothrus badius 235 
momota, Momotus 63, 234 
Momotus momota 63, 234 
monacha, Pachycephala 304 
monachus, Myiopsitta 126, 233 
Monasa nigrifrons 234 
mondetoura, Claravis 227, 233 
montagnii, Penelope 233 
montanus, Passer 36 
montivagus, Aeronautes 198, 234 
moresbyi, Lonchura atricapilla 4 
Morphnus guianensis 27—30 
Motacilla flava 310 
motmot, Ortalis, 196, 198 
mulsant, Acestrura 234 
munda, Serpophaga 234 
muraria, Tichodroma 145-6 
murina, Phaeomyias 234 
Muscicapa griseisticta 305 
Muscipipra vetula 79 
Muscisaxicola maculirostris 230, 234 
musicus, Turdus 140 
Myadestes ralloides 235 
mycteria, Jabiru 160 
Myiagra inquieta 310 
Myiarchus sp. 234 

— cephalotes 234 

— ferox 234 


XIV 


— swainsoni 234 

— tyrannulus 230, 234 
Myioborus brunniceps 232, 235 

— castaneocapillus 198 
Myiodynastes chrysocephalus 234 

— maculatus 234 
Myiopagis gaimardii 234 

— viridicata 234 
Myiophobus fasciatus 64, 234 
Myiopsitta monachus 126, 233 
Myiozetetes similis 234 
Myrmothera simplex 198 
Myrmotherula longicauda 234 
mystacea, Aplonis 308 
mystaceus, Platyrinchus 64 


nana, Grallaricula 254 

— , Sylvia 301 
Nannopsittaca panychlora 198 
nasutus, Tockus 82 
nattereri, Anthus 286, 289, 290 

— , Hylopezus 280, 290 
naumanni, Falco 4 
nematura, Lochmias 198 
Neochmia phaeton 308 
Neomixis striatigula 8 
Neopelma aurifrons 125 
nevermanni, Lonchura 310 
Newtonia sp. 8 
nigra, Astrapia 46 

— , Coracopsis 159 
nigricans, Sayornis 234 
nigriceps, Eremopterix 301 
nigricollis, Sporophila 275, 279, 289 
nigrifrons, Monasa 234 
nigrocapillus, Phyllomyias 252 
nigrolineata, Ciccaba 63 
noctua, Strix 137 
nordmanni, Glareola 4 
notata, Coturnicops 78 
Nothocercus bonapartei 248 
Nothoprocta pentlandii 233 
Notiochelidon cyanoleuca 234 
— flavipes 254 
noveboracensis, Seiurus 77 
Nyctibius aethereus 171 
— griseus 234 
Nycticorax nycticorax 233 
Nyctidromus albicollis 234 
Nyctiphrynus ocellatus 171 
Nystalus chacuru 234 

— maculatus 234 


obscura, Elaenia 234 

— , Lonchura atricapilla 4 

=) oO arise 235 
obsoletum, Camptostoma 234 
obsoletus, Crypturellus 233 

— , Salpinctes 229 
occipitalis, Phylloscopus 186 
Oceanodroma hornbyi 159 

— markhami 159 


ocellatus, Nyctiphrynus 171 
ochrocephala, Amazona 205-9 
Ochthoeca cinnamomeiventris 250, 252 
— leucophrys 234 
odomae, Metallura 254 
Odontorchilus branickii 71 
Oenanthe deserti 301 
— hispanica 142 
— isabellina 82 
olivaceus, Vireo 235 
olivater, Turdus 198 
olivetorum, Hippolais 4 
ophthalmicus, Chlorospingus 235 
— , Phylloscartes 234 
Opisthoprora euryptera 254 
oratrix, Amazona 205-9 


oratrix hondurensis, Amazona subsp. 


nov. 205-9 
Oreophasis derbianus 16 
Oreopsar bolivianus 235 
Oreotrochilus adela 234 
Oriolus oriolus 82, 140 
ornatus, Spizaetus 168 
Ortalis guttata 233 

— motmot 196, 198 
Orthotomus sutorius 213 
oryzivora, Scaphidura 235 
Otus albogularis 253 

— capnodes 125 

— choliba 228, 234 

— hoyi 228, 234 


Pachycephala monacha 304 
Pachyramphus castaneus 234 

— major 273 

— polychopterus 234, 273 

— versicolor 250 

— viridis 234 
pacificus, Apus 35 
pallatangae, Elaenia 198 
palmarum, Dendroica 76 

— , Thraupis 235, 287° 
paludicola, Riparia 213 
palustris, Sporophila 78, 174, 288, 290 
panychlora, Nannopsittaca 198 
Panyptila cayannensis 234 
papa, Sarcoramphus 233 
papuensis, Collocalia 304 
Paradigalla sp. 119 

— brevicauda 38-47 

— carunculata 38-47 

— intermedia 39-47 
Paradisaea apoda 141, 309 

— minor 119 

— raggiana 309 
paraguaiae, Gallinago 198 
Parisoma boehmi 244 

— buryi 244 

— layardi 244-7 

— lugens 244 

— subcaeruleum 244 
Parotia sefilata46 


XV 


Parula pitiayumi 235 
Parus ater 114—5 
Passer iagoensis 118, 156 

— montanus 36 
Passerella iliaca 101-5 
Pastor senex 67-8 
Patagona gigas 234 
pectorale, Dicaeum 305 
pectoralis, Polystictus 127 
pelegrinoides, Falco 149 
peli, Scotopelia 4 
Penelope jacquacu 233 

— montagnii 233 
pennatus, Hieraaetus 316 
pentlandii, Nothoprocta 233 
peregrinus, Falco 32, 156, 233 
personatus, Atlapetes 198 
perspicillata, Pulsatrix 234 
peruviana, Tangara 316-8 
Phacellodomus striaticeps 234 
Phaeomyias murina 234 
Phaethon aethereus 157 
Phaethornis pretrei 234 

— superciliosus 234 
phaeton, Neochmia 308 
Phalacrocorax brasilianus 233 
Phalcoboenus australis 242 

— megalopterus 233 
phasianellus, Dromococcyx 170 
phelpsi, Cypseloides 196, 198 
Pheucticus aureoventris 232, 235 
philbyi, Alectoris 82 
philomelos, Turdus 160, 312-3 
philippensis, Bubo 34 
philippinus, Merops 213 
phoenicius, Tachyphonus 196, 198 
Phoenicopterus minor 4 

— ruber 4 
Phonygammus diamondi 130-1 

— purpureoviolaceus 130-1 
Phylidor rufus 234 
Phyilomyias burmeisteri 234 

— nigrocapillus 252 

— sclateri 234 
Phylloscartes ophthalmicus 234 

— sylviolus 172 

— ventralis 234 
Phylloscopus affinis 186 

— armandii 186 

— collybita 160 

— davisoni 186 

— emeiensis 180 

— griseolus 186 

— occipitalis 186 

— proregulus177—93 

— reguloides 186 

— schwarzi 186 

— sibilatrix 180 

— subaffinis 186 
Phytotoma rutila 234 
Piaya cayana 234 
Pica pica 140 


picina, Mearnsia 35 
Picoides lignarius 228, 234 
picta, Chloephaga 241 
picui, Columbina 233 
Piculus chrysochloros 234 

— rubiginosus 198, 234 
Picumnus cirratus 234 
picumnus, Dendrocolaptes 234 
Picus canus 127 
pileata, Pionopsitta 78, 262 
Pinarornis plumosus 4 
Pinguinus impennis 159 
Pionopsitta pileata 78, 262 
Pionus sp. 234 

— maximiliani 234 

— menstruus 234 

— seniloides 126 

— tumultuosus 126, 258 
Pipile jacutinga 78 

— pipile 233 
Pipraeidea melanonota 231, 235 
Pipreola arcuata 252 

— lubomirskii 72-3 

— pulchra 73 

— riefferii 72 
Piranga flava 235 
Pitangus sulphuratus 234 
pitiayumi, Parula 235 
Pitohui incerta 308 
placens, Poecilodryas 308 
plancus, Polyborus 233 
plantensis, Embernagra 232, 235 
platyrhynchum, Electron 234 
Platyrinchus leucoryphus 64 

— mystaceus 64 
Plegadis falcinellus 309 
Ploceus galbula 82 
plumbea, Columba 233 

—  , Ictinia 233 
plumosus, Pinarornis 4 
Podiceps dominicus 198 
Poecilodryas placens 308 
Poecilurus scutatus 234 
poecilurus, Knipolegus 197-8 
poliocephalus, Accipiter 306 
poliogaster, Accipiter 279 
Polioptila dumicola 232, 234 
Polyborus plancus 233 


polychopterus, Pachyramphus 234, 273 


polyosoma, Buteo 233, 258 
Polystictus pectoralis 127 
Polytmus milleri 198 
pomarina, Aquila 149 
pompadora, Treron 213 
Poospiza boliviana 235 

— lateralis 281, 290 

— melanoleuca 235 

— torquata 232, 235 

— whitii 235 
Porzana fusca 33 
prasinus, Aulacorhynchus 250 
preciosa, Tangara 316 


Premnoplex adusta 198 
pretiosa, Claravis 279 
pretrei, Amazona 282, 290 
— , Phaethornis 234 
Prinia flaviventris 212 
— hodgsonii 213 
— inornata 213 
promeropirhynchus, Xiphocolaptes 234, 
260 


Promerops cafer 245 

— gurneyi 245 
proregulus, Phylloscopus 177—93 
przewalski, Grallaria 72 
Psarocolius angustifrons 235 

— atrovirens 235 

— decumanus 235 
Pseudochloroptila pseudochloroptila 245 

— symonsi 245 
Pseudocolaptes boissonneautii 258 
Psittaculirostris desmarestii 307 
Psittrichas fulgidus 307 
ptaritepui, Crypturellus 195, 198 
Pterocles coronatus 301 

— lichtensteini 301 

— senegallus 301 
Pterodroma sp. 118 

— feae 129, 157 
Pteroglossus castanotis 234 
pucherani, Campylorhamphus 255 
Puffinus spp. 118 

— assimilis 118, 157 

— lherminieri 118 
pulchra, Pipreola 73 
Pulsatrix koeniswaldiana 282 

— perspicillata 234 
punctatus, Thamnophilus 234 
punctulata, Lonchura 36 
purpurea, Ardea 156 
purpureoviolaceus, Phonygammus 130-1 
Pycnonotus blanfordi 213 

— finlaysoni 213 

— goiavier 36 

— virens 75 
pygargus, Circus 4 
Pyriglena leuconota 234 
Pyrocephalus rubinus 234 
Pyrrhomyias cinnamomea 234, 258 
pyrrhophia, Cranioleuca 234 
pyrrhops, Hapalopsittaca 253, 257—62 
pyrrhopterus, Lycocorax 119 
Pyrrhura molinae 233 


querquedula, Anas 309 


radiatus, Carpococcyx 126 
raggiana, Paradisaea 309 
ralloides, Myadestes 235 
Ramphastos toco 234 
— tucanus 234 - 
Ramphocelus carbo 173, 235 
Ramphotrigon megacephala 172, 220-3 
razae, Alauda 118, 156-7 


regalis, Heliangelus 69 
reguloides, Phylloscopus 186 
Regulus ignicapillus 160 
Rhabdornis inornatus 36 
Rhamphomantis megarhynchus 307 
Rhea americana 78 
Rhinomyias albigularis 128 
gularis 128 

Rhynchotus rufescens 61-2 
ridibundus, Larus 239 
riefferii, Chlorornis 251 

, Pipreola 72 

Riparia paludicola 213 

risora, Alectrurus 64, 78 
rixosus, Machetornis 234 
roraimae, Automolus 198 
ruber, Phoenicopterus 4 
rubescens, Anthus 120 
rubetra, Saxicola 213 
rubidiceps, Chloephaga 241 
rubiginosus, Piculus 198, 234 
rubinus, Pyrocephalus 234 
rubricollis, Campephilus 234 

, Drymophila 285, 290 
rubrogenys, Ara 227, 233 
rufa, Casiornis 234 

, Formicivora 63 
rufescens, Acrocephalus 48—50 
, Rhynchotus 61-2 

, Sericornis 46 
ruficapilla, Vermivora 273 
ruficapillus, Thamnophilus 234, 285 
ruficollis, Corvus 301 

, Sporophila 79, 174 

, Tachybaptus 32 
rufigularis, Falco 233 
rufiventris, Lurocalis 251, 253 
, Turdus 235 
rufocarpalis, Calicalicus sp.nov. 5-10 
rufosuperciliata, Syndactyla 230, 234, 285 
rufulus, Anthus 213 

, Troglodytes 198 

rufus, Caprimulgus 234 

, Furnarinus 232, 234 

, Phylidor 234 

, Tachyphonus 231, 235 
ruspoli, Tauraco 11-5 
russatum, Todirostrum 198 
rustica, Hirundo 36 

rutila, Phytotoma 234 
rutilans, Xenops 234 

rutilus, Cypseloides 234, 272-3 
Rynchops flavirostris 4 


Salpinctes obsoletus 229 
Saltator aurantiirostris 65, 235 
maximus 235 
samarensis, Eurylaimus 126 
sapphirina, Hylocharis 283-4 
Sappho sparganura 234 
Sarcoramphus papa 233 
Satrapa icterophrys 234 


XV1l 


saularis, Copsychus 213 
Saxicola caprata 213, 304 
jerdoni 210-8 

rubetra 213 

sayaca, Thraupis 232, 235 
Sayornis nigricans 234 
Scaphidura oryzivora 235 
scheepmakeri, Goura 307 
Schistochlamys melanopis 235 


schomburgku, Micropygia 275, 277, 289 


schulzi, Dryocopus 79 

schwarzi, Phylloscopus 186 
sclateri, Phyllomyias 234 
scolopacea, Eudynamys 34 
Scotopelia peli 4 

scutatus, Poecilurus 234 

sefilata, Parotia 46 

Seiurus noveboracensis 77 
selimbauensis, Lonchura atricapilla 4 
seloputo, Strix 34 

semifasciata, Tityra 275, 278, 289 
Semioptera wallacei 141 
semipalmatus, Anseranas 309 
semitorquatus, Arremon 294-8 
senegalensis, Ephippiorhynchus 4, 57 
senegallus, Pterocles 301 

senex, Heterornis 67-8 

, Pastor 67-8 

, sturnus 67 

, Temenuchus 67 
seniloides, Pionus 126 

sericeus, Sturnus 68 
sericocaudatus, Caprimulgus 171 
Sericornis rufescens 46 
Sericossypha albocristata 255 
Sericulus aureus 309 

Serinus canaria 180 

Serpophaga cinerea 234 

munda 234 

serranus, Turdus 251 
serrirostris, Colibri 234 

severa, Ara 233 

sibilatrix, Phylloscopus 180 
Sicalis flaveola 235 

luteola 274 

sidamoensis, Heteromirafra 15, 75 
similis, Myiozetetes 234 
simplex, Myrmothera 198 
sinensis, Centropus 213 

Sitta canadensis 140 

whiteheadi 140 

Sittasomus griseicapillus 234 
sittoides, Diglossa 235 

smithii, Hirundo 213 

solitarius, Harpyhaliaetus 233 

, Tinamus 78 

sordida, Thlypopsis 235 
sparganura, Sappho 234 
sparverius, Falco 233 

speciosa, Columba 170 
speciosum, Conirostrum 231, 235 
Spizaetus ornatus 168 


— tyrannus 169 
Spizastur melanoleucus 168 
Spizella breweri 130 

— taverneri 130 
Sporophila caerulescens 235 

— cinnamomea 174, 288, 290 

— falcirostris 173, 287 

— hypochroma 174 

— nigricollis 275, 279, 289 

— palustris 78, 174, 288, 290 

— ruficollis 79, 174 
squamiceps, Turdoides 82 
Steatornis caripensis 196, 198 
steeril, Eurylaimus 126 
Stephanophorus diadematus 287, 290 
Sterna fuscata 81 

— hirundo 33 

— sumatrana 33 
stictolopha, Lophornis 235-6 
Stigmatura budytoides 232, 234 
stoltzmanni, Urothraupis 252 
Streptopelia chinensis 34 
Streptoprocne zonaris 196, 198, 234 
stresemanni, Zavattariornis 15 
striata, Dendroica 77 

— , Geopelia 34, 304 

— sumatrensis, Lonchura 4 
striaticeps, Phacellodomus 234 
striatigula, Neomixis 8 
striativentris, Melanocharis 306 
striatus, Butorides 82 
ls alba 137 

aluco 34 

— huhula 283 

— hylophila 283 

— noctua 137 

— seloputo 34 
Struthio camelus 4 
Sturnus albofrontatus 68 

— senex 67 

—  sericeus 68 
stygia, Lonchura 310 
subaffinis, Phylloscopus 186 
subcaeruleum, Parisoma 244 
Sublegatus modestus 232, 234 
Suiriri suiriri 232, 234 
Sula dactylatra 81 

— leucogaster 81, 157 
sulphuratus, Pitangus 234 
sulphurea, Cacatua 126 
sulphurescens, Tolmomyias 234 
sumatrana, Sterna 33 
sumatrensis, Lonchura striata 4 
superciliaris, Leptogon 234 
superciliosus, Accipiter 279 

— , Phaethornis 234 
sutorius, Orthotomus 213 
swainsoni, Buteo 281, 290, 315-6 

— , Myiarchus 234 
sylvatica, Turnix 33 
Sylvia sp. 244 

— nana 301 


XV1ll 


sylviolus, Phylloscartes 172 
symonsi, Pseudochloroptila 245 


- Synallaxis azarae 234 


— chinchipensis 125 
— frontalis 234 


Syndactyla rufosuperciliata 230, 234, 285 


Tachybaptus ruficollis 32 
Tachyeres brachydactyla 242 
Tachyphonus phoenicius 197-8 

— rufus 231, 235 
taciturnus, Arremon 294-8 
talpacoti, Columbina 233 
‘Tangara cabanisi 16-8 

— cyanoptera 198 

— peruviana 316-8 

— preciosa 316 
Tanysiptera galatea 307 

— hydrocharis 307 
‘Taphrospilus hypostictus 234 
Taraba major 234 
tataupa, Crypturellus 233 
‘Tauraco leucotis 11-5 

— ruspoli 11-5 
taverni, Spizella 130 
Temenuchus albofrontatus 67 

— senex 67 
‘Terathopius ecaudatus 4, 82 
Tersina viridis 235 
thalassinus, Colibri 234 
Thalurania furcata 234 
Tharinephalis sp. 234, 285 

aroyae 234 

— caerulescens 234 

— doliatus 234 

— insignis 198 

— punctatus 234 

— ruficapillus 234, 285 

— torquatus 285 
Thamnornis chloropetoides 8 
Theristicus caudatus 232, 233 
Thlypopsis sordida 235 
Thraupis bonariensis 235 

— palmarum 235, 287 

— sayaca 232, 235 
Thripophaga berlepschi 71 
Thryothorus genibarbis 234 
thula, Egretta 233 
Tiaris fuliginosa 175, 288 

— obscura 235 
Tichodroma muraria 145-6 
Tigrisoma fasciatum 233 
Tyuca atra 243 
timoriensis, Megalurus 304, 310 
Tinamus major 233 

— solitarius 78 
tinnunculus, Falco 149, 156 
Tityra cayana 278 

— inquisitor 278 

— semifasciata 275, 278, 289 
tobaci, Amazilia 197-8 
Tockus bradfieldi 4 


— nasutus 82 
toco, Ramphastos 234 
Todirostrum cinereum 234 

— russatum 198 
Tolmomyias sulphurescens 234 
torquata, Ceryle 234 

— , Melanopareia 172, 286, 289 

— , Poospiza 232, 235 
torquatus, Atlaptes 235 

— , Buarremon 251 

— , Thamnophilus 285 
Treron pompadora 213 
Trichoglossus haematodus 127 
Tringa glareola 33 
trochilirostris, Campylorhamphus 234 
Troglodytes aedon 234 

— cobbi 242 

— rufulus 198 
Trogon collaris 234 

— curucui 234 
tucanus, Ramphastos 234 
tumultuosus, Pionus 126, 258 
turcosa, Cyanolyca 250-1, 258 
Turdoides squamiceps 82 
Turdus amaurochalinus 232, 235 

— chiguanco 235 

— fuscater 235, 258 

— ignobilis 197-8 

— iliacus 140 

— merula 112-4 

— musicus 140 

— olivater 198 

— philomelos 160, 312-3 

— rufiventris 235 

— serranus 251 
Turnix sp. 33 

— sylvatica 33 
tyrannulus, Myiarchus 230, 234 
Tyrannus melancholicus 234 
tyrannus, Spizaetus 169 
Tyto alba 156, 242 


undulatus, Crypturellus 233 
uncinatus, Chondrohierax 62, 168, 233 
uniformis, Chloropipo 198 

unirufa, Cinnycerthia 252 

Upucerthia harterti 229, 234 

Upupa epops 19-26, 82 

Uropsalis lyra 234 

Urothraupis stoltzmanni 252 
urubitinga, Buteogallus 195, 198 


vana, Lonchura 46 
Vanellus duvaucelii 213 
vanikorensis, Collocalia 35 
varia, Mniotilta 77 


X1X 


vasa, Coracopsis 159 
vauxi, Chaetura 272 
Veniliornis sp. 234 

— frontalis 234 

— fumigatus 234 
ventralis, Phylloscartes 234 
Vermivora ruficapilla 273 
verreauxi, Leptotila 233 
versicolor, Amazilia 197-8 

— Pachyramphus 250 
versicolorus, Brotogeris 227, 233 
vespertinus, Falco 4 
vetula, Muscipipra 79 
vinacea, Amazona 282 
vinaceigula, Egretta 4 


virens amadoni, Andropadus nom.nov. 


re 

— , Pycnonotus 75 
Vireo leucophrys 251 

— olivaceus 235 
virgata, Ciccaba 171 
virginianus, Bubo 234 
viridicata, Myiopagis 234 
viridis, Anurolimnas 275, 276, 289 

— , Carpococcyx 126 

— , Pachyramphus 234 

— , Tersina 235 
Volatinia jacarina 235 
Vultur gryphus 233 


waldeni, Aceros 127-8 
wallacei, Semioptera 141 
wetmorei, Buthraupis 255 
whiteheadi, Sitta 140 
whitii, Poospiza 235 
williami, Metallura 254 
wolfi, Aramides 160 


xanthogastra, Carduelis 235 
xanthogonys, Heliodoxa 196, 198 
Xenops rutilans 234 
Xenopsaris albinucha 65, 231, 234 
Xenospingus concolor 314 
Xiphocolaptes major 234 

— promeropirhynchus 234, 260 
Xiphorhynchus guttatus 234 
ypiranganus, Emberizoides 289, 290 


Zavattariornis stresemanni 15 
Zenaida auriculata 233 
Zimmerius bolivianus 234 

— chrysops 73 

— cinereicapillus 73 
zonaris, Streptoprocne 196, 198, 234 
Zonotrichia capensis 198, 235, 358 


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156 
57 
157 
170 
230 
242 
243 
243 


305 
311 


Line 9 
Line 25 
Line 47 
Line 48 
Line 49 
Line 50 
Line 50 


- Line 19 


Line 22 
Line 42 
Line 7 
Line 18 
Line 41 
Line 45-49 


Line 4 
Line 44 


XX 
CORRECTIONS TO TEXT 


Rynchops not Ryncops 

sulphurea not sulphurata 

Ardea bourne: not Ardea purpurea bournet 

Falco neglectus not Falco tinnunculus neglectus 

Falco alexandri not Falco tinnunculus alexandri 

Falco madens not Falco peregrinus madens 

Tyto detorta not Tyto alba detorta 

Calonectris edwardsu not Calonectris diomedea edwardsiu 
Puffinus boydi not Puffinus assimilis boydi 

Dromococcyx not Dromoccocyx 

dorbignyi not dorbygni 

Phalcoboenus not Phalcobaenus 

cristata not cristat 

See for Erratum to Prof. Richard Chander’s article in Vol. 117 
page 158. 

gigantea not lugens 

P. Gregory’s present address: PO Box 168, Katherine 0851, 
Northern Territory, Australia. 


Contents Vol. 117 No. 4 line 23 philomelos not Philomelos 


Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


OL 
671 
B63 
6B/RD 


ISSN 0007-1595 


Bulletin of the 


British Ornithologists’ Club 


Edited by 
Dr D. W. SNOW 


Volume 117 No. 1 


March 1997 


FORTHCOMING MEETINGS 


Tuesday 20 May 1997. ANNUAL GENERAL MEETING AT 6 p.m., followed 
by a Club Social Evening. There will be no speaker, but Members are invited to 
bring along one or two siides (or a specimen!) of a bird of topical interest, and to 
speak for not more than 5-10 minutes about it. The aim will be to generate 
discussion, and to facilitate the exchange of information between Members. 

Applications to the Hon. Secretary by 6 May, please, including subjects to be raised, 
and any special facilities required. 


Tuesday 17 June 1997. Robin Woods will speak on the “Atlas of Breeding 
Birds of the Falkland Islands” which was published in early 1997. Having been 
obsessed with Falkland Islands ornithology for over 40 years, Robin has produced 
the two popular books on the birds of the Falklands, in 1975 and 1982, and the 
Guide to Birds of the Falkland Islands (1988), which gave full treatment to all species 
recorded. He was the first to prove, in 1960, the breeding of the Great Shearwater in 
the Falklands, and to outline the importance of mature, ungrazed tussac grass to 
about 30 breeding species. His latest book is the result of an extensive ten-year 
fieldwork and census project, begun in 1983/84. 

Applications to the Hon. Secretary by 3 Fune, please. 


Tuesday 15 July 1997. Tim Appleton will give a talk titled “From Greenfields 
to Ramsar’. Tim is well-known to all visitors to Rutland Water, having been - 
Reserves Manager there for over 21 years. Arriving with the experience of working 
at Slimbridge, he was involved in the design and planning stages for the creation of 
this highly successful project, and has managed this reserve ever since; he has also 
been co-organiser of the British Birdwatch Fair, which enjoys a worldwide 
reputation. 

Applications to the Hon. Secretary by 1 Fuly, please. 


Meetings are held in the Sherfield Building of Imperial College, South 
Kensington, London SW7, at 6.15 p.m. for 7 p.m. The nearest Tube station is at 
South Kensington, and car parking facilities are available; a map of the area will be 
sent to members, on request. The cash bar is open from 6.15, and a buffet supper, of 
two courses followed by coffee, is served from about 7.00. (A vegetarian menu can be 
arranged if ordered at the time of booking.) Informal talks are given on completion. 


Overseas Members visiting Britain are particularly welcome at meetings. 
For details in advance, please contact the Hon. Secretary, Cdr M. B. 
Casement, OBE, RN, Dene Cottage, West Harting, Petersfield, Hants. 
GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1997 


Apart from single copies made for the purposes of research or private study, or 
criticism or review, as permitted under UK law, no part of this publication may be 
reproduced, stored or transmitted in any form or by any means, except with prior 
permission in writing of the publishers, or in accordance with the terms of licences 
issued by the Copyright Licensing Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; 
for address see inside back cover. 


1 Bull. B.O.C. 1997 117(1) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 117 No. 1 Published 21 March 1997 


REPORT OF THE COMMITTEE FOR 1996 


Meetings. Eight evening meetings were held in 1996, six in the 
Ante-room, and two in the Senior Common Room of the Sherfield 
Building at Imperial College, London. A total of 286 (201 members and 
85 guests) attended these meetings, which represented an average 
attendance of 36. The programme of speakers during the year again 
covered a wide variety of ornithological subjects, and was presented by 
speakers both from Britain and from overseas. A ninth meeting, in the 
form of a social evening on 10 December, was cancelled due to a clash of 
dates. The 860th meeting of the Club was celebrated with an excursion 
to Selborne on Friday 31 May, attended by 17 members and 9 guests. 

In addition, a symposium on “Avian taxonomy from Linnaeus to 
DNA” was held jointly with the Linnean Society, at Burlington House, 
Piccadilly, on Saturday 23 March. The total number attending, 
including speakers, was 101. 

Committee. The Committee met eight times during the year, and the 
attendance was 78%. The Committee accepted the ruling from the Charity 
Commissioners that the Hon. Editor should cease to be a Trustee of the 
Club, if he was to receive an honorarium for his work, and the Rules of the 
Club were accordingly amended and approved at the AGM on 21 May. 

Terms of reference were agreed for a sub-committee to coordinate 
the sponsorship, production and sales of the Club publications. The 
Committee is greatly indebted to Mrs Moore for her continuing work 
as Chairman of the Publications Sub-Committee (see below), and also 
to Dr R. P. Prys-Jones and Dr R. A. Cheke for their time and expertise 
as editors for these works. 

The Trustees of the Herbert Stevens Trust Fund met with the Com- 
mittee on two occasions to give advice on the performance of the Fund. 
The Committee is most grateful to Mr Nigel Crocker, Mr Richard Price 
and Mr Peter Oliver for their time and expertise in this matter. 

Owing to increasing pressures of other work, Mr John Farnsworth 
tendered his resignation as Hon. Treasurer, in July, to take effect as 
soon as a replacement could be found. The committee is grateful to Mr 
David Montier for agreeing to take over as Acting Hon. Treasurer. He 
is now responsible for the financial aspects, whilst membership and 
other aspects of the post are being progressively delegated to the Hon. 
Secretary, and other members of the Committee. 

It is with great regret that the Committee reports the deaths in the 
past year of the following six members: Dr D. R. Aspinwall 
(1981-1996), Dr G. C. Ellsworthy (1985-1996), Mr K. W. Henshall 
(1986-1996), Mr J. L. McKean (1978-1996), Mr J. G. Parker 
(1977-1996 and Committee Member 1979-1983), and Sir William 
Wilkinson (1987-1996). 


Report of Committee 2 Bull. B.O.C. 1997 117(1) 


Publications Sub-committee. Publication of Occasional Publi- 
cations No. 2 (Manuscripts and Drawings in the Ornithology and 
Rothschild Libraries of the Natural History Museum at Tring, authored 
by Mrs F. E. Warr) has been completed; and production of No. 3 
(Avian Egg-shells: an Atlas of Scanning Electron Micrographs, by 
Konstatin Mikhailov) is well advanced for publication in 1997. Work 
on a Bird Atlas of Uganda is proceeding according to plan, with the aim 
of publication in mid-late 1998, subject to finalisation of a Licence 
Agreement with the authors, and adequate sponsorship funding. 

Membership. There were 567 paid-up Members at 31 December 
1996—322 with addresses in the U.K., and 245 overseas. Active 
recruitment from the newly-elected Members of the Union continued, 
resulting in 38 new Members joining the Club during the year. 

Bulletin. Volume 116 consisted of 272 pages, and contained 39 main 
papers and 9 shorter (In Brief) contributions. Two new species were 
described, both illustrated with colour plates: a new munia from Borneo, 
and a new pipit from South Africa, the latter especially remarkable as 
coming from an ornithologically well known area. Four new subspecies 
were described, and an account of some very little known birds from the 
Solomon Islands included field notes and figures of two forms that almost 
certainly represent undescribed taxa. New distributional data were pub- 
lished for Cameroon, Tanzania, Brazil, Ecuador, Bolivia, Solomon 
Islands and South Georgia. Among papers of special interest may be 
mentioned the description and analysis of hybrid hummingbirds of the 
genus Phaethornis; a review of the migrations of the Arctic Tern, includ- 
ing overland records and ocean records collected over nearly 50 years by 
the Royal Naval Birdwatching Society; a survey of geographical variation 
in sexual dichromatism in birds; and the first record of the breeding of 
Forbes’s Plover in Central Africa. There were also substantial papers 
presenting new data on the status and distribution of little known Andean 
birds, and a fully annotated checklist of the birds of South Georgia. The 
70 authors were from 15 countries and 6 continents. 

Once again, the Committee is greatly indebted to Mary Muller for her 
compilation of the annual index of authors, contents and scientific names. 

Bulletin Sales. Sales to non-member subscribers totalled 142, with 
16 in the U.K. and 126 overseas (31 countries). Of the latter, 50 
addressees were in the United States of America, and 16 in Germany. 

Finance. Most of the income of the Club continues to come from 
subscriptions, both from members and non-member subscribers, and 
from investment income. During the year 1996, the Trustees of the 
Herbert Stevens Trust Fund, with the approval of the Committee, 
switched the investments of the Fund away from a portfolio of 
individual fixed interest and equity stocks into three unit trust funds 
designed specifically for charities. Apart from obtaining a wider spread 
of risk, this should allow the Club to receive income without tax being 
deducted, rather than having to make separate claims to recover the tax 
from the Inland Revenue after the end of the tax year. 

The main expenditure related to the printing and distribution of the 
Bulletin, and is well in excess of the subscription income. The increase 
in subscription incomes applicable from 1 January 1997 will go some 


Annual General Meeting 3 Bull. B.O.C. 1997 117(1) 


way towards reducing this imbalance. One major non-recurring 
expense during the year was the binding of the Chairman’s set of copies 
of the Bulletin at a cost of £840. 

Accounts for 1996, which are not yet available, will be tabled at the 
Annual General Meeting, and published subsequently in the Bulletin. 


ANNUAL GENERAL MEETING 


The Annual General Meeting of the British Ornithologists’ Club will 
be held in the Ante-room of the Sherfield Building, Imperial College, 
London SW7 at 6.00 p.m. on Tuesday 20 May 1997. 


AGENDA 
1. Minutes of the 1996 Annual General Meeting (see Bull. Brit. Orn. 
Cl. 116(4): 201-202). 

Report of the Committee and Accounts for 1996. 

The Bulletin. 

The election of Officers. The Committee proposes that: 

(i) The Reverend T. W. Gladwin be elected Chairman, wice 
Mr D. Griffin, who remains as a member of the Committee. 

(ii) Mrs A. M. Moore be elected as Vice-Chairman vice The 
Reverend T. W. Gladwin. 

(iii) Mr D. J. Montier (currently Acting Hon. Treasurer) be 
elected as Honorary Treasurer, vice Mr S. J. Farnsworth, who 
retires. 

(iv) Commander M. B. Casement, OBE, RN, be re-elected Hon. 
Secretary. 

(v) Mr N. J. Redman be elected a member of the Committee vice 
Mr R. E. F. Peal who retires by rotation, and is ineligible for 
re-election. 

(vi) Professor C. J. Feare be appointed Hon. Editor, in succession 
to Dr. D. W. Snow, with effect from about September 1997. 

5. Any other business of which notice shall have been given in 

accordance with Rule (12). 


oon 


By Order of the Committee 
MICHEAL CASEMENT, Honorary Secretary 


The eight hundred and sixty-fourth meeting of the Club was held on Tuesday 5 
November 1996, at 6.15 p.m. 29 Members and 13 guests attended. 

Members present were: D. GRIFFIN (Chairman), N. D. HUNTER (Speaker), M. A. 
Apcock, Miss H. Baker, J. W. BarrRINGTON, B. H. Beck, I. R. BisHop, Mrs D. M. 
BraD.ey, P. BuLL, D. R. CALDER, Cdr M. B. CaseEMENT RN, Professor R. J. CHANDLER, 
Dr R. A. CuHexe, Dr R. A. F. Cox, S. J. FARNSwortu, F. M. GAunTLETT, Rev. T. W. 
GLapwin, C. A. R. Heim, B. A. E. Marr, D. J. Montier, Mrs A. M. Moore, Mrs 
M. N. MUuLLeER, P. J. O_tver, Dr R. P. Prys-Jones, N. J. RepMAN, P. J. SELLAR, Dr 
N. J. Secvar, S. A. H. StatuaM, N. H. F. STONE. 

Guests attending were: Mrs J. HUNTER, Mrs S. GrirFin, Miss G. BonuHaM, R. GILBEY, 
Mrs J. C. BuLL, Mrs J. B. CaLper, Mrs C. R. CASEMENT, Mrs M. H. Gaunt Lett, Mrs 
J. M. Giapwin, Mrs M. Montier, P. J. Moore, Mrs M. Ouiver, M. PALING. 

After dinner, Nigel Hunter spoke on ‘“The Birds of Botswana’’, illustrated by an 
impressive selection of photographic slides taken by Brian Bushell, coupled with species 
distribution maps reproduced from Huw Penry’s 1994 Bird Atlas of Botswana. 


Editorial note 4 Bull. B.O.C. 1997 117(1) 


His talk focused on providing an overview of the significance of Botswana’s birds in the 
Sub-Saharan context. After a brief geographical and ecological background, he pointed 
out that Botswana has no “‘endemics”’ as such, but the Slaty Egret Egretta vinaceigula is 
currently known only to breed in the Okavango Delta. Botswana is also important for the 
presence of four regional endemics—Cape Vulture Gyps coprotheres, Bradfield’s Hornbill 
Tockus bradfieldi, Short-clawed Lark Certhilauda chuana, and Boulder Chat Pinarornis 
plumosus. In addition, there are four species with a “‘special status’”’ because of concern 
about their status elsewhere—White-backed Night Heron Gorsachius leuconotus, African 
Skimmer Ryncops flavirostris, Pel’s Fishing Owl Scotopelia peli and Yellow-billed 
Oxpecker Buphagus africanus. 

Due to its relatively low human poulation and general unsuitability for arable 
agriculture, Botswana still contains significant populations of the larger species, 
increasingly threatened elsewhere. Examples include Ostrich Struthio camelus (as a 
free-range wild species), Saddle-billed Stork Ephippiorhynchus senegalensis, both 
Flamingoes Phoenicopterus ruber and P. minor, seven birds of prey including the Bateleur 
Terathopius ecaudatus, Wattled Crane Bugeranus carunculatus, Kori Bustard Ardeotis kort, 
and Ground Hornbill Bucorvus leadbeateri. 

Concerning migrant species, he suggested that African migrants were not especially 
significant in a Botswana context, though relatively little was yet known about their 
movements in the region during the African winter (i.e. May—September). But Botswana 
is important as a wintering area (October—April) for Palearctic migrants from Europe, 
and especially for 12 of the 28 species under this category. These are White Stork Ciconia 
ciconia, Montagu’s and Pallid Harrier Circus pygargus and C. macrourus, both Red-footed 
Kestrels Falco amurensis and F. vespertinus, Lesser Kestrel F. naumanni, Black-winged 
Pratincole Glareola nordmanni, \cterine and Olive-tree Warbler Hippolais icterina and 
H. olivetorum, River Warbler Locustella fluviatilis, and Lesser Grey and Red-backed 
Shrikes Lanius minor and L. collurio. 

Although the Okavango Delta usually receives most attention, the Kalahari region of 
Botswana comprises by far the greater proportion of the country. He illustrated the 
ornithological flavour of this ecosystem with slides of some 50 typically representative 
species. The strong influence rain can play in a semi-arid system, prone to drought, was 
shown by slides of 27 additional species which deploy real opportunism by taking 
short-term advantage of the periodic emergence of surface water, usually in “pans’’. 


Proposed additions to the genus Lonchura: Addenda and 
Corrigenda 

On p. 154 of the paper in Bull. Brit. Orn. Cl. 115(3), details of the 
type of Lonchura atricapilla obscura subsp. nov. were not included. The 
two specimens that were at hand when the taxon was named were 
AMNH specimens 447931 and 447932, both males, collected at Parit, 
Tjempaga, Sampit, S. Borneo, 3 July 1935. The first of these (447931) 
should have been designated as the holotype. 

The endings of three of the new names proposed contravene the rules 
laid down in the International Code of Zoological Nomenclature (1985 
edition), and should be amended as follows: p. 148, amend bzgilalae 
to bigilale: (see Article 31 of the Code); p. 149, amend moresbyae to 
moresbyi (see Article 31); p. 155, amend selimbaue to selimbauensis (see 
Appendix D, 1v, of the Code). 

References to Paynter & Storer (1970, on p. 140 and subsequently, 
should be to Paynter & Storer 1968 (Peters’ Check-list, vol. 13 not 
vol. 14). On p. 142, the name Lonchura striata sumatrensis was proposed 
not by Chasen & Hoogerwerf (1941) but by Chasen (1939, Treubia 17, 
p. 183), who designated as its type a specimen in the Buitenzorg 
Museum, no. 11249. K 

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Plate 1. (upper) Photograph of presumed male Calicalicus rufocarpalis taken by C.A.D. 
on 1 March 1991 approximately 22 km southeast of Toliara by road (17 km by air), on 
the old track to St Augustin and less than 10 km east of La Table (the type locality of 
C. rufocarpalis). (lower) Photograph of male Calicalicus madagascariensis taken by T. S. 
Schulenberg in early March 1992 near Ifaty, 32 km north of Toliara. 


S. M. Goodman et al. 5 Bull, BOC; £997 117(1) 


A new species of vanga (Vangidae, Calicalicus) 
from southwestern Madagascar 


by Steven M. Goodman, A. F. A. Hawkins & 
Charles A. Domergue 


Received 19 March 1996 


The Red-tailed Vanga Calicalicus madagascariensis (Linnaeus, 1766) is 
the smallest member of the family Vangidae, all of which are forest 
species and endemic to the Malagasy faunal region (Madagascar and 
the Comoros). C. madagascariensis, the only previously named species 
of this genus, is an arboreal gleaning insectivore. Red-tailed Vangas 
often forage on the outer parts of canopy trees and shrubs on 
medium-sized insects such as caterpillars and crickets (Rand 1936, 
Langrand 1990, Yamagishi et al. 1992, Eguchi et al. 1993). 

All members of the Vangidae except the Red-tailed Vanga and Blue 
Vanga Cyanolanius madagascarinus are either limited to one of the three 
major Malagasy biogeographic regions (Eastern, Western and 
Southern) or have populations divided into subspecies that are 
restricted to one or two of these regions (Rand 1936, Appert 1968, 
1970, Langrand 1990). Thus in general their distributions seem closely 
linked with vegetational communities. 

The Red-tailed Vanga is widely distributed on Madagascar (Fig. 1), 
though it appears to be absent from a few areas in the northwest 
(Hawkins 1994) and the extreme south and southwest (Langrand 1990), 
and it occurs patchily elsewhere in its range (Appert 1968, 1970). No 
geographic variation has been previously recognized in this species. It 
is found in all native forest types and in degraded and secondary areas 
adjacent to primary forest (Rand 1936, Langrand 1990, Hawkins 1994). 

In mid-June 1948 Philippe Milon collected two Calicalicus near 
Tuléar (Toliara) in the extreme southwestern corner of the island. 
After comparison of these two specimens to material of Calicalicus 
madagascariensis from across the island held in the American Museum 
of Natural History (AMNH), the Natural History Museum (BMNH), 
the Field Museum of Natural History (FMNH), and Muséum National 
d’Histoire Naturelle (MNHWN), Paris, we have concluded that they 
represent a distinct and previously undescribed species. Here we 
propose to call this new species 


Calicalicus rufocarpalis sp. nov. 


Holotype. Female, Muséum National d’Histoire Naturelle, Paris, 
1974.510, Tuléar [=Toliara], Province de ‘Toliara, Madagascar, 
collected 19 June 1948, by Colonel Philippe Milon; field number PM 
4396. The specimen had a granular ovary, measuring 5 X 2 mm, and 
with small “‘microscopique”’ egg follicles. 

Further details about the specimen, not written on the museum 
ticket, are recorded in Milon’s field catalogue (MNHN). Soft part 


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S.M. Goodman et al. 7 Bull. B.O.C. 1997 117(1) 


) TABLE 1 
Measurements (in mm) and mass (g) of Calicalicus madagascariensis and C. rufocarpalis 
bill length 

wing exposed from anterior 

chord tail culmen edge of nostril tarsus weight 
rufocarpalis nov. sp. 
MNHN 1974.5102 = 64 57 11:9 2d | 23.6 17.4 
MNHN 1974.5082 62 57 11.6 8.5 24.7 16.1 
mean 63.0 57.0 11.8 8 24.2 16.8 


madagascariensis’ 67.4, 1.88 48.4, 2.04 10.5, 0.52 7.8, 0.40 20.4, 0.83 15.5, 1.5 
65-72 45-54 9.4-11.6 7.0—-8.7 18.5-22.4 14.0-18.0 
(n=53) (n=52) (n=53) (n=52) (n=43) (n=6) 


statistical F=10.9 F=34.5 F=10.3 F=12.1 F=38.8 F=1.1 
comparison 
(ANOVA) p=0.002 p=0.0001 p=0.002 p=0.0001 p=0.0001 p=0.34 


‘Sexes combined. Presented as mean, standard deviation, minimum-maximum (number 
of specimens). 


colours: iris lemon with a slight tawny wash, beak brownish-black with 
greyish-blue near base. Weight: 17.4 g. The bird was collected near La 
Table [approximately 20km southeast of oliara], which is here 
designated as the type locality. An analysis of its stomach contents 
conducted by R. Paulian identified one cerambycid beetle and a large 
grasshopper, almost certainly Locusta. 

Diagnosis. Calicalicus rufocarpalis is distinguished from C. madagas- 
cariensis by differences in external measurements and plumage 
characters. Colour names and numbers are after Smithe (1975). 

The sample of C. madagascariensis (n=53) showed no sexual 
dimorphism in external measurements and in all analyses presented 
herein sexes of adult specimens are combined. C. rufocarpalis has 
distinctly shorter wings and longer tail, bill and tarsus than C. 
madagascariensis (‘Table 1). In all cases there is no to little overlap in 
measurements between these two species and the differences are all 
statistically significant. There is considerable overlap in weight. 

The crown and mantle of female rufocarpalis are generally a 
continuous near Brownish-olive (29) or Olive (30), while in 
madagascariensis the Plumbeous (78) crown contrasts sharply with the 
Brownish-olive or Olive mantle. The upper wing-coverts of rufocar- 
palis are distinctly Brick-red (132a) which contrasts with the 
Brownish-olive wing-coverts with Clay-color (26) fringes in female C. 
madagascariensis. The Brick-red pigment on the tail in rufocarpalis 
tends to be restricted to the area bordering the central shaft and absent 
from the tip, while in madagascariensis all the tail feathers, except the 
central pair, are almost entirely Brick-red. No specimen of male 
C. rufocarpalis is known (see Discussion section). 


S. M. Goodman et al. 8 Bull. BLOvO P99’ 18K 1) 


Measurements of the holotype (mm). Wing (chord) 64, tail 57, exposed 
culmen 11.9, bill length from anterior edge of nostril 9.1, tarsus 23.6. 

Paratype. Female, Muséum National d’Histoire Naturelle, Paris, 
1974.508, Tuléar (Toliara), Province de Toliara, Madagascar, collected 
17 June 1948, by Colonel Philippe Milon; field number PM 4391 (see 
‘Table 1 for measurements). 

Other information on this specimen was found in Milon’s field 
catalogue (MNHN). The specimen was collected near a stone quarry at 
La Table, 13 km southeast of 'Toliara, and was part of a mixed species 
foraging flock composed of four or five Thamnornis chloropetoides, four 
or five Newtonia, and one Neomixis striatigula. The iris was light 
yellowish-brown and the bill brownish-black with a slightly bluish 
wash, and greyish-blue at the base. The bird weighed 16.1 g. 

Etymology. ‘The name rufocarpalis is derived from the Latin and 
refers to the distinct Brick-red wing converts of females of the new 
species. 

English name. We suggest Red-shouldered Vanga. 

Range. Only known from ‘Toliara region of southwestern 
Madagascar. 

Specimens examined. C.. rufocarpalis, 222, Tuléar (Toliara) (MNHN). 
C. madagascariensis, 6334, 5299, 3 unsexed: Anaborano 1g (AMNH); 
Andapa 12 (AMNH); Bejofo 1¢, 12 (AMNH); Bezona 1¢ (AMNH); 
Col Pierre Radama 1g (AMNH); Eminiminy 1¢ (AMNH); Fanovana 
1¢ (BMNH); Fito 233, 292 (AMNH), 1¢ (MNHN); Forét de 
Vohibasia 1g (FMNH); Forét Sihanaka 4992 (BMNH); Fort Dauphin 
(Tolagnaro) 14, 12 (MNHN); Iampasika 12 (AMNH), 104, 29° 
(BMNH), 12 (MNHN); Ivohibe 23.4, 229 (AMNH), 13 (BMNH), 12 
(MNHN); Lac Iotry 12 (AMNH); Manombo 23g, 299° (BMNH), 14, 
12 (MNHN); Maroantsetra region 11¢¢, 699, 2 unsexed (AMNH), 
355, 229 (MNHN); Maromandia 1g (AMNH), 13, 12 (MNHN); 
Marotony 2¢¢, 12 (AMNH); Montagne d’Ambre 2¢¢, 12 (MNHN); 
Nosy Be 3¢¢ (MNHN); Périnet 2¢¢, 12 (MNHN); Sakaraha 19° 
(MNHN); Tabiky 2¢¢, 12 (AMNH), 2¢¢ (BMNH), 12 (MNHN); 
Tsarakibany 12 (MNHN); Tsiandro 4¢¢, 622 (AMNH), 434, 12, 1 
unsexed (BMNH), 235, 292 (MNHN); Vondrozo 443, 622 (AMNH), 
24d, 12 (BMNH), 1¢ (MNHN). 


Discussion 


The only known material of C. rufocarpalis is the holotype and 
paratype, both females. In both cases, they have Brick-red 
upperwing-converts, a character in C. madagascariensis restricted to 
males (Langrand 1990). Milon, the collector of both rufocarpalis 
specimens, was clearly aware of the unusual circumstances of a female 
Calicalicus having Brick-red wing-coverts. On the original specimen 
label, of the holotype, with hand writing in the distinct style of Milon, 
he underlined the female sex symbol and wrote “ov. [aire] g. [ranulaire] 
assez important,’ thus removing any doubt associated with the proper 
sexing of the specimen. No mention was made of the two Toliara birds 
in his book on Malagasy birds (Milon et al. 1973). In examination of 


S. M. Goodman et al. 9 Bull,» BOC A997 IAF (A) 


C. madagascariensis specimens we found a few sub-adult males in 
female plumage. In all cases, males seem to obtain simultaneously 
Brick-red wing-coverts and other aspects of the adult male plumage 
(e.g. black throat feathers). Examples of males in this plumage include 
individuals taken at Tabiky on 17 November 1929 (AMNH 413072), at 
Bezona, near Ambanja, on 25 November 1930 (AMNH 413078), and 
at Vondrozo on 6 July 1929 (BMNH 1931.8.18.1516). 

Calicalicus rufocarpalis is only known from the Toliara region, while 
C. madagascariensis occurs in much of the remaining forested areas of 
the island (Langrand 1990). The nearest known sites to La Table for 
C. madagascariensis are 110 km northeast of Toliara near Sakaraha 
(MNHN 1973.553), 59 km northeast of Sakaraha in the Forét de 
Vohibasia (FMNH 380006), and 32 km north of Toliara near Ifaty 
(photographed by T. S. Schulenberg). These sites are shown in 
Figure 1. 

A male Calicalicus attending a nest was photographed by C.A.D. on 
1 March 1991, approximately 22 km southeast of Toliara by road 
(17 km by air), on the old track to St Augustin. The site (23°26'S, 
43°48’E) was located about 500m south of the turn-off towards 
Ambohimahavelona, which lies on the Onilahy River, and less than 
10km east of La Table. This individual had a plumage pattern 
different from typical C. madagascariensis (Plate 1). The most 
noticeable differences of the male in the photograph are the completely 
rufous upperwing-coverts as well as rufous on the mantle and lower 
nape as compared to typical C. madagascariensis in which the rufous 
feathers are restricted to the marginal, lesser and some median 
wing-coverts, and the mantle and upperparts are grey. The 
photographed bird has a yellow iris and C. madagascariensis possesses a 
dark brown iris. Further, the bird also has a distinctly paler tail and 
more heavily washed rufous underparts than C. madagascariensis. 
Given the combination of characters, in particular the iris colour, 
we conclude that the bird photographed by C.A.D. is referable to 
C. rufocarpalis. 

Given the limited number of records of Calicalicus from the 
southwest, it is unclear if C. madagascariensis and C. rufocarpalis 
are sympatric. The closest known sites of their occurrence are Ifaty 
(C. madagascariensis) and Toliara (C. rufocarpalis) which were until 
recently part of the same forest block, separated only by the seasonal 
Fiherenana River. Further research is needed to clarify the distribution 
of these two species in southwestern Madagascar. 

C. rufocarpalis has a shorter wing and tail and longer bill and tarsus 
than C. madagascariensis. The forest stature of the spiny bush in 
the Toliara region is distinctly shorter than that of the dry deciduous 
forest and humid forest, the latter types being the domain of 
C. madagascariensis. The differences in the measurements of the two 
species are presumably related to these ecological variables. 

The current status of C. rufocarpalis is unknown. With the exception 
of the photo from southeast of Toliara, we are unaware of any recent 
records of it from the region between the Fiherenana River (north of 
Toliara and south of Ifaty) south to the Onilahy River (Fig. 1). Within 


S. M. Goodman et al. 10 Bull BLOVELT99SY VEAt 


a few kilometres of the Toliara city centre there still remains some 
forested areas, although generally degraded. Further, the forests of 
southwestern Madagascar are under heavy human pressure associated 
with exploitation of trees for charcoal and timber. Recent fieldwork in 
the region of Toliara and St Augustin has failed to reveal further 
records of C. rufocarpalis and it is presumed to be rare or sparsely 
distributed. We strongly suspect that C. rufocarpalis has a broader 
geographic range than currently known. It might be a species 
associated with the coastal plain and plateau of the calcareous Mahafaly 
Plateau, which runs from near Toliara south to Androka. This region of 
Madagascar is ornithologically poorly known and has not been the 
focus of any thorough inventory. Other species of vertebrates, such as 
the recently described carnivore Galidictis grandidieri (Wozencraft 
1986, Goodman 1996), are known to have a parallel geographic 
distribution. 


Acknowledgements 


We are most grateful to curators of various museums who made material available 
for study: G. Barrowclough and M. LeCroy, AMNH; M. Walters, BMNH; J.-F. and 
C. Voisin, MNHN. Goodman’s visit to the American Museum of Natural History was 
made possible by a grant from the Frank M. Chapman Fund. T. Schulenberg kindly 
made available photographs. We are grateful to Olivier Langrand and Tom Schulenberg 
for comments on an earlier version of this MS. 


References: 

Appert, O. 1968. La répartition géographique des vangidés dans la région du Mangoky et 
la question de leur présence aux différentes époques de l’année. Ozsseau 38: 6-19. 

Appert, O. 1970. Zur Biologie der Vangawiurger (Vangidae) Sudwest-Madagaskars. Orn. 
Beob. 67: 101-133. 

Eguchi, K., Yamagishi, S. & Randrianasolo, V. 1993. The composition and foraging 
behaviour of mixed species flocks of forest-living birds in Madagascar. Ibis 134: 
91-96. 

Goodman, S. M. 1996. A subfossil record of Galidictis grandidieri (Herpestidae: 
Galidiinae) from southwestern Madagascar. Mammalia 60: 150-151. 

Hawkins, A. F. A. 1994. Forest degradation and the western Malagasy forest bird 
community. Ph.D. thesis, University of London. 

Langrand, O. 1990. Guide to the Birds of Madagascar. Yale Univ. Press. 

Milon, P., Petter, J.-J. & Randrianasolo, G. 1973. Oiseaux. XXXV. Faune de 
Madagascar. ORSTOM & CNRS, Tananarive and Paris. 

Rand, A. L. 1936. Distribution and habits of Madagascar birds. Bull. Amer. Mus. Nat. 
Hist. 72: 143-499. 

Smithe, F. B. 1975. Naturalist’s Color Guide. American Museum of Natural History, 
New York. 

Wozencraft, W. C. 1986. A new species of striped mongoose from Madagascar: 
J. Mammalogy 67: 561-571. 

Yamagishi, S., Eguchi, K., Nakamura, M., Nagata, H. & Randrianasolo, V. 1992. 
Comparative foraging ecology of Vangids (Vangidae). Pp. 39-45 in: S. Yamagishi 
(ed.), Social structure of Madagascar higher vertebrates in relation to their adaptive 
radiation. Osaka City University, Osaka, Japan. 


Address: Steven M. Goodman, Field Museum of Natural History, Roosevelt Road at 
Lake Shore Drive, Chicago, Illinois 60605, U.S.A. and World Wide Fund for 
Nature, Aires Protégées, BP 738, Antananarivo (101), Madagascar. A. F. A. 
Hawkins, BP 8511, Antananarivo (101), Madagascar. Charles A. Domergue, Institut 
Pasteur, BP 1274, Antananarivo (101), Madagascar. 


© British Ornithologists’ Club 1997 


L. Borghesio 11 Bull. B.O.C. 1997 117(1) 


Observations on the ecology of Tauraco 
ruspolu and T. leucotis in southern Ethiopia 


by L. Borghesio 
Received 28 December 1995 


Two species of the genus Tauraco are endemic to the Abyssinian 
plateau: Prince Ruspoli’s Turaco T. ruspolit and the White-cheeked 
Turaco T. leucotis, the latter with two subspecies, the nominate and 
donaldson. ‘They form a superspecies with T. hartlaubi of the Kenyan 
plateau. JT. leucotis is widely distributed across most of Ethiopia, and 
even penetrates marginally into Eritrea and south-eastern Sudan, while 
T. ruspoli has a very restricted distribution in the southern part of the 
country (Fig. 1). 

Following the theory first proposed by Moreau (1958), the ancestor 
of the two turacos was cut into two by the Rift Valley during a dry 
period, giving rise to leucotis on the western and ruspoli on the eastern 
side. When the climate ameliorated, Jleucotis crossed the Rift, 
differentiated there into the subspecies donaldsoni, and pushed ruspoli 
southwards. Still later, leuwcotis crossed the barrier again, driving 
ruspoliu into its last refuge on the southern margin of the highlands. 
According to this view, TJ. ruspolit would be a relict species, 
competitively inferior to T. /eucotis and gradually disappearing. Owing 
to this and to habitat alteration within its extremely restricted range, 
T. ruspolu is therefore now considered an endangered species (Ash & 
Gullick 1989, Dellelegn 1991, Collar et al. 1994) and is listed in the 
African Red Data Book (Collar & Stuart 1985). 

In order to evaluate the status of T. ruspolii, a survey was undertaken 
between 23 March and 5 June 1995. Some data on its habitat 
requirements and those of T. leuwcotis were collected, providing new 
insight into the competition between and evolution of the two species. 
This new information is reported in this paper. 

The area surveyed extends over about 35,000 km* in the Ethiopian 
administrative regions of Sidamo, Borana and Bale, and lies at the 
southern margin of the Abyssinian plateau. Elevation reaches over 
2000 m in the northern sections, descending southwards to about 
800 m. Habitats comprise Podocarpus gracilior montane forests in the 
wetter areas (mainly in the north); in drier localities, but still receiving 
a good amount of water, often in the form of mist, another kind of 
montane forest develops, dominated by Juniperus procera. The forests 
gradually merge into increasingly dry Acacia woodlands where 
elevation and rainfall decrease. 

During the survey, 49 sites were visited (Fig. 2) within and around 
the known range of T. ruspolii. The species was found in all the 
previously known localities, including Genale (5°40'N, 30°32'E), where 
it was thought to have disappeared in recent times (Ash & Gullick 
1989), and in numerous previously unreported sites. The habitats 
frequented ranged from Podocarpus and Juniperus forests and forest 


2 


L. Borghesio 12 Bull. B.O.C. 1997 117(1) 


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Figure 1. Distribution of the Ethiopian turacos. 


margins up to Acacia-dominated woodlands, in the latter only as 
long as the preferred food plants, especially figs (Ficus sycomorus, 
F. thonningu, F. vasta), were present. Densities in Podocarpus forests 
were very low and only four individuals were met with there 
during over 90 hours of search in different localities; since the fruits of 
isolated Podocarpus trees growing outside forests were readily eaten by 
T. ruspolii, Podocarpus forests should not in themselves represent a 
hostile environment for the species; in fact the main reason for its 
absence is probably related to competition from T. leucotis (see below). 
In summary, the preferred habitats of T. ruspolii were forest margins 
and woodlands, where 10 to 20 individuals were regularly observed in a 
single day. Also Juniperus forests, the habitat where the species was 
initially re-found by Benson (1942, 1945), held good numbers. 


L. Borghesio 13 Bull. B.O.C. 1997 117(1) 


6°N 


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Probable range of T.ruspolii 


) S limit of Podocarpus forest and patches 
Figure 2. Map of the survey area. 


Figure 3 shows the altitudinal distribution of the observations of 
T. ruspoli, which ranged from 1250 to 1860 m a.s.l. The upper limit 
roughly coincides with that of the most elevated localities in the study 
area; the lower one, which is reached after descending steeply from 
about 1500 m, appears to form an altitudinal limit to the species’ 
distribution. As can be seen in Figure 2, the localities where T. ruspoli 
occurred were all within, or very near to, the 1500 m contour line. 

The geographic distribution of T. ruspoliu, inferred from present 
information, is recorded in Figure 2. In this map the northern limit of 
the species is seen to coincide with the southern limit of Podocarpus 


L. Borghesio 14 Bull. B.O.C. 1997 117(1) 


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1300 1400 1500 1600 1700 1800 1900 


elevation (m a.s.l.) 


Figure 3. Altitudinal distribution of observations of Tauraco ruspolii. 


forests, marked by a shaded line in the map, while elsewhere the range 
approximately follows the 1500 m contour line. This results in a surface 
area extending over about 7740 km”. It was suggested by Collar & 
Stuart (1985) that the range of the species could extend outside the area 
represented in Figure 2, towards the town of Ginir (7°08’N, 40°43’E), 
in a northeastward direction; but this seems unlikely, as it would 
imply a range extension of over 150 km in an area mostly lying below 
1300 m. 

T.. leucotis was also observed in the study area (Fig. 2). In contrast to 
T.. ruspolu, highest densities occurred in Podocarpus. ‘The two species 
often occurred within a few kilometres of each other, but they were 
found together in the same habitat in only three places (Fig. 2): at 
Wadera (5°48'N, 39°20’E), in a Podocarpus forest where leucotis greatly 
outnumbered ruspolii, at Zembaba (5°53’N, 39°10’E), and at Bore 
(5°38'N, 38°48’E), both in Podocarpus forest margins. Within the range 
of ruspoli, leucotis was never found in woodlands, and it was less 
common in Podocarpus forest margins; thus, it was almost exclusively 
present in Podocarpus forests. Erard & Prévost (1970) found that at 
Wadera ruspoli occurred inside and leucotis outside the forest, but 
this was not the case in 1995. Outside the range of ruspolz, leucotis 
occurred commonly in habitats drier than forests (i.e. forest margins, 
riverine vegetation and woodlands) that were similar to those occupied 
by ruspoli alone in the area of their co-existence. Elsewhere in 
Ethiopia, J. leucotis is not considered a specialised species of 
Podocarpus forests, but is seen to exploit a relatively wider range of 
habitats than its congener (Urban & Brown 1971, pers. obs.), and is 
reported to occur at elevations as low as 850 m in the Omo Valley (Fry 
et al. 1988). 


L. Borghesio 15 Bull. B.O.C. 1997 117(1) 


The present observations suggest that the two Ethiopian turacos are 
largely separated by habitat in the areas where they come in contact. 
This was particularly evident in some localities, such as Wadera and 
Sede (5°31'N, 39°08’E) where Podocarpus forest abutted onto mixed 
Acacia-conifer woodland. There the two species replaced each other 
along the habitat transition, with little or no overlap. This pattern, in 
which different species replace each other very sharply in different 
habitats, is common in the Musophagidae (Fry et al. 1988, Dillingham 
& Moreau 1961). 

Since their habitat overlap is restricted, the hypothesis of strong 
interaction between the two species no longer seems to hold. T. ruspoli 
is probably prevented from extending its range into the Podocarpus 
forest by the presence of the related species, but since in sympatric 
situations J. lewcotis occupies a more restricted range of habitats than it 
does in allopatry, competition is not likely to be a severe threat for 
ruspoli, which seems to be better adapted than its relative to its 
non-forest habitat. As there is no evidence that the range of T. ruspoliu 
is presently declining owing to competition, it is also unlikely that this 
happened in the past; hence the species should probably not be 
considered a gradually disappearing relict as was proposed by Moreau 
(1958), but rather as a stenoecious species showing preference for 
middle altitude habitats of intermediate dryness between montane 
forest and thorn-bush. 

In the light of these conclusions, the conservation status of 7. ruspoli 
should probably be considered in a less dramatic way than it has been 
in the past. The range of the species is very small, but competition is 
clearly not a threat to its survival, and, even more important than this, 
T. ruspolu is clearly not dependent on forests. Its preferred habitats, 
owing to their greater dryness, usually have to stand a much lower 
pressure from the expanding human population, and are not as severely 
threatened as are most Ethiopian forests. 

Three other species of birds, also with very restricted altitudinal 
distributions, are found in the same area in the southern Ethiopian 
highlands (Ethiopian Bush-crow Zavattariornis stresemanni, White- 
tailed Swallow Hirundo megaensis, Sidamo Long-clawed Lark Hetero- 
mirafra sidamoensis). ‘Taking this into consideration, it would seem that 
this region in the past, probably during a dry period, became separated 
from the rest of the Ethiopian highlands by the barriers formed by the 
Rift Valley to the west and by the huge block of the Bale mountains 
(reaching over 4200 m; Fig. 1) to the north; this isolation provided the 
conditions for the evolution of the four restricted-range bird species. 

T. ruspolii seems therefore to have evolved in situ in the southern 
highlands of Ethiopia and, owing to its restricted altitudinal 
distribution, which certainly reduces its chances of dispersion, it may 
have never occupied a much wider range than today. On the other hand 
T. leucotis, which, as Moreau (1958) correctly pointed out, was initially 
restricted to the western side of the Rift valley, thanks to its ability 
to colonise localities at lower elevations, was able to cross the barrier 
and occupy a much wider area, which at present almost completely 
encircles T. ruspolit. 


H. Gomez de Silva G. 16 Bull. B.O.C99F 17 @) 


Acknowledgements 


Many persons gave their help and their friendship to me during the execution of this 
work in Ethiopia and in Italy. They are listed here in alphabetical order: Dr John Ash, 
Mr John Atkins, Ato Yilma Dellelegn, Prof. Mauro Fasola, Ato Amare Kifle, Dr 
Yalemtsehay Mekonnen, Ato Tilaye Nigussie, Shemseddi Nuri, Mr Giampiero Pagani, 
Prof. Anacleto Sabbadin, Mr Per Ole Syvertsen, Ato Abebe Takwa, Ato Akale Yemane, 
Ato Melaku Zendu. The research was supported by a grant from the Bird Exploration 
Fund, and received assistance from the Ethiopian Natural History and Wildlife Society 
and the Instituto Italiano di Cultura of Addis Ababa. 


References: 

Ash, J. S. & Gullick, T. M. 1989. The present situation regarding the endemic breeding 
birds of Ethiopia. Scopus 13: 90-96. 

Benson, C. W. 1942. A new species and ten new races from southern Abyssinia. Bull. 
Brit. Orn. Cl. 63: 8-19. 

Benson, C. W. 1945. Notes on the birds of southern Abyssinia. [bis 87: 489-509. 

Collar, N., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to Watch 2. The World List of 
Threatened Birds. BirdLife Conservation Series no. 4. ICBP, Cambridge. 

Collar, N. & Stuart, S. 1985. Threatened Birds of Africa and Related Islands. The 
ICBP/IUCN Red Data Book Part 1. 3rd edn. ICBP/IUCN, Cambridge. 

Dellelegn, Y. 1991. Prince Ruspoli’s Turaco. Walia 13: 29-35. 

Dillingham, I. H. & Moreau, R. E. 1961. Musophagidae in northwestern Tanganyika 
territory. Ibis 103: 294-295. 

Erard, C. & Prévost, J. 1970. New facts on the distribution of Tauraco ruspolii Salvadori. 
Bull. Brit. Orn. Cl. 90: 157. 

Fry, C. H., Keith, S. & Urban, E. K. 1988. The Birds of Africa. Vol. 3. Academic Press. 

Moreau, R. E. 1958. Some aspects of the Musophagidae. Jbis 100: 67-112. 

Urban, E. K. & Brown, L. H. 1971. A Checklist of the Birds of Ethiopia. Addis Ababa 
Univ. Press, Addis Ababa. 


Address: L. Borghesio, C. Re Umberto 42, 10128 Torino, Italy. 


© British Ornithologists’ Club 1997 


Further observations on the nesting of the 
Azure-rumped ‘T'anager 


by Héctor Gomez de Silva Garza 
Received 15 April 1995 


This paper describes observations on a nest of the Azure-rumped 
Tanager Tangara cabanisi in the El] Triunfo Biosphere Reserve, 
southeastern Chiapas, Mexico. The nest was discovered by Angie 
Tyner on 26 April 1993, while the birds were still building it, and from 
1 to 17 May the author was able to make occasional visits while 
conducting observations on the natural history of the Horned Guan 
Oreophasis derbianus and censusing the birds in the upper montane 
forest of El Triunfo. 


H. Gomez de Silva G. 17 Bull. B.O.C. 1997 117(1) 
TABLE 1 


Data on nest sites of Tangara cabanisi; measurements in metres. All nests were at a point 
where the branch forked 


Height of Height of Distance from Distance 


tree nest edge of crown to trunk Branch 
Data from Long & ——115.4-36.1 9.7—32.5 0.5-2 3-14 horizontal 
Heath (1994) 
1993 nest c. 25 15.5 1.5-2 c.4 horizontal 


The Azure-rumped Tanager is a little known species with an 
extremely small range (Collar et al. 1992). Long & Heath (1994) 
presented the most detailed account of the breeding biology of this 
species, including data from 9 nests discovered between 1983 and 1990. 
They concluded that Azure-rumped ‘Tanagers breed between 
mid-April and mid-June, nest in trees over 15m tall with “a wide 
canopy of spreading branches’’, and place their nests “‘in the top half of 
the tree, some distance from the main trunk, approaching the end of a 
long-horizontal branch, at a point where the branch forked.”’ 

Not surprisingly, the 1993 nest fits this description well (Table 1). A 
noteworthy difference, however, is that the nest was not in the “humid 
evergreen broadleaf forest’’ that is the tanagers’ typical habitat (and 
where all of Long & Heath’s nests were found), but in a pine tree in the 
adjacent “‘Cupressus-Pinus community” at higher elevations (habitats as 
described in Long & Heath 1991). Heath & Long (1991: 227) reported 
that they occasionally saw the tanager up to 1700 m above sea level, 
where some plant species from the humid evergreen broadleaf forest 
follow the river valleys up into the lower slopes of the conifer belt. The 
1993 nest was very near such a valley, and indeed the incubating bird 
nearly always flew toward the valley to forage, but it is significant that 
the bird searched for similar characteristics for a nest site even within 
vegetation of very different structure and composition from its typical 
habitat. This suggests that the choice of the above-mentioned nest-site 
characteristics may be a fixed “‘species-specific’’ pattern. 

The average duration of incubation recesses (8.00 min, s.d. 4.12, 
n=45) was slightly higher than the averages for two nests (6.7, s.d. 3.4; 
5.7, s.d. 3.2) reported in Long & Heath (1994). This is not surprising, 
since the 1993 nest was not surrounded on all sides by the proper 
habitat and the birds would have been expected to require longer 
periods of time to find enough of the fruits which form the main part of 
their diet. 

Long & Heath (1994) reported that the incubation period is ‘“‘around 
14 days’’. It is typically 13-14 days in the genus Tangara (Isler & Isler 
1987). Unfortunately, the bird may have already been incubating for 
one or more days by the time that my observations of the 1993 nest 
began, but the young had not yet hatched by 13.00 hr on the 15th day 
of observation; they had already hatched by 13.00 hr of day 16. The 


H. Gomez de Silva G. 18 Bull. B.O.C. 1997 117(1) 


incubation period, therefore, was at least 15 days. Long & Heath (1994) 
report that the incubation constancies for two nests observed were 73.9 
and 81.7%, calculated using the formula of Skutch (1976). From my 
data, I calculated a constancy of 67.4%. The range falls within the 
60-80% of daylight hours reported for most tanagers (Skutch 1989) but 
is lower than that recorded in the previous Azure-rumped Tanager 
nests, probably owing to the longer average duration of recesses. It is 
important to note that it was necessary to modify Skutch’s formula for 
calculating incubation constancy owing to the different number of 
incubation recesses and sessions observed in full (45 and 35); the 
average duration was used instead of the sums of all recesses and 
sessions. This calculation includes very brief recesses of 3 min (when a 
Cathartes vulture soared low overhead), 1 min (in order to catch an 
insect in flight) and a few seconds (in order to chase a pair of 
Blue-hooded Euphonias Euphonia elegantissima which began to build 
their nest in an adjacent tree). 

Eggs of the genus Tangara are usually ‘“‘whitish (sometimes tinged 
with colour), and speckled brown or lavender, especially about the large 
end”’ (Isler & Isler 1987). The eggs of the Azure-rumped Tanager were 
previously unknown, but in the 1993 nest could be seen to be whitish, 
with a very pale pink wash and heavy red-brown speckling. 


Acknowledgements 


These observations were carried out while conducting research with Fernando 
Gonzalez-Garcia for the Instituto de Ecologia. Angie Tyner discovered the nest and 
kindly told me of its location; Al DeMartini, Alain Huc and Leobardo Pérez helped while 
I watched the nest. I would also like to thank for their companionship Marco Tulio 
Argueta, Ismael Galvez, Laura Noble, Rafael Solis, Sofia Solérzano and the Watts 
family. 


References: 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
T. A. III & Wege, D. C. 1992. Threatened Birds of the Americas: the ICBP/IUCN 
Red Data Book. International Council for Bird Preservation. Cambridge, U.K. 

Heath, M. & Long, A. 1991. Habitat, distribution and status of the Azure-rumped 
Tanager Tangara cabanisi in Mexico. Bird Conservation International 1: 223-254. 

Isler, M. L. & Isler, P. R. 1987. The Tanagers: natural history, distribution and 
identification. Smithsonian Institution Press. 

Long, A. & Heath, M. 1991. Flora of the El Triunfo Biosphere Reserve, Chiapas, 
Mexico: a preliminary floristic inventory and the plant communities of Polygon I. 
Anal. Instit. Biol. UNAM, Mexico, Ser. Bot. 62(2): 133-172. 

Long, A. & Heath, M. 1994. Nesting ecology and helping behavior in the Azure-rumped 
Tanager in Mexico. Condor 96: 1095-1099. 

Skutch, A. F. 1976. Parent Birds and Their Young. Univ. Texas Press. 

Skutch, A. F. 1989. Life of the Tanager. Cornell Univ. Press. 


Address: Héctor Gémez de Silva G., Centro de Ecologia, UNAM, Apartado Postal 
70-275, C.P. 04510 Cuidad Universitaria, México, D.F., Mexico. 


© British Ornithologists’ Club 1997 


P.G. P. Ericson 19 Bull. B.O.C. 1997 117(1) 


Swedish records of the eastern Palearctic 
Hoopoe subspecies Upupa epops saturata 


by P. G. P. Ericson 
Received 20 January 1996 


Southern Sweden is at the northwestern extreme of the European 
breeding distribution of the Hoopoe Upupa epops. 'The Hoopoe breeds 
in Sweden in very low numbers, mainly on Oland (Pettersson 1994). 
Besides this small breeding population, a significant number of 
vagrant individuals are observed outside the breeding areas, most often 
in spring and autumn (SOF 1990). Very rarely, Hoopoes winter in 
Sweden. 

It is commonly assumed that all Hoopoes breeding or migrating 
through Sweden belong to the European subspecies Upupa epops epops 
(SOF 1990), of which Sweden is actually the type locality. U. e. epops is 
a typically pale subspecies inhabiting Europe, northwestern Africa, 
northwest India, Sinkiang in China, and Russia east to the Ob—Yenisey 
watershed (Cramp 1985). Most of the other subspecies occurring in, or 
near, the Palearctic are very similar to U. e. epops, differing only in 
being smaller and deeper rufous or duller in coloration with a longer 
bill. The different subspecies also have varying amounts of white on the 
wing and crest feathers. Only the east Palearctic subspecies U. e. 
saturata difters from U. e. epops in being distinctly darker, especially on 
the mantle and breast (Lo6nnberg 1909, Vaurle 1959). Except for a 
possible intergrading zone between nominate epops and saturata in the 
upper Yenisey watershed and in Tibet (Cramp 1985), there is no 
tendency towards darker birds anywhere within the breeding range of 
U. e. epops. 

This paper evaluates the hypothesis that the subspecies U. e. saturata 
occurs regularly in Sweden. The work was initiated by the finding of 
a very dark individual in the province Blekinge in southern Sweden 
in December 1993, which obviously tried to winter there (Stromberg 
1994). This bird is similar to other Swedish specimens in_ the 
collections of the Swedish Museum of Natural History which closely 
match the description of U. e. saturata. The difference in coloration of 
typical U. e. epops and typical U. e. saturata is obvious and subspecific 
determinations can fairly easily be done in the hand, and probably also 
in the field. However, as in most cases of this kind of geographical 
variation, individuals exhibiting intermediate characters occur. In such 
cases the subspecific allocation is often dependent on the personal 
judgement of the observer. With the application of modern optical 
techniques to quantify colours, this subjectivity can be reduced. 


Materials and methods 


A total of 17 adult specimens of Upupa e. epops and 16 of U. e. saturata 
(including the type and two paratypes) were studied. All specimens 


P.G. P. Ericson 20 Bull. B.O.C. 1997 117(1) 


(listed in Appendix) are kept in the collections of the Swedish Museum 
of Natural History (NRM). 

The mantle and breast of each individual were illuminated by 
tungsten-halogen light through a fibre-optic probe and the reflectance 
spectra were obtained by an Ocean Optics S1000 diode array 
spectrometer. From the spectrogram, the CIELAB (CIE 1986) colour 
parameters a (red-green scale), b (yellow—blue scale) and L (grey scale) 
were calculated. a, 6 and L are orthogonal axes producing a 
three-dimensional colour space designed to fit the human perception of 
colour hues. The degree to which the measured colour was serrated, 
the ‘‘colourfulness”, was estimated by calculating chroma as (a*+b*)'/” 
In a bivariate plot of a and b, the hues can be expressed as the angle 
of the vector from the origin to the coordinates (a', b'). The hues of 
different individuals, expressed as arctan (b/a), were ‘then compared. 

Ideally, for fully comparable results readings should be taken at 
homologous points of each individual in order, but this goal is very 
difficult to achieve in practice. Early in the work it proved that the 
repeatability of the readings was low, and that it is critical to 
standardise both the angle between the probe and the surface, and 
the pressure with which it is attached. In order to minimise the 
intra-observer variation, nine readings were taken from each specimen 
and the median value was used in the statistical analyses. 

Statistical difference between samples were tested by a two-tailed 
Student’s f-test. 


Results 


Sexual dimorphism 

The difference between the sexes in plumage coloration was assessed 
in order to determine whether the sexes could be pooled in the analyses 
of the difference between the two subspecies. Although there is no 
obvious way to determine the sex of a Hoopoe in the field, sexual 
dimorphism involving general size (significant) and colour of throat and 
breast (slight) has been reported in the subspecies U. e. epops (Cramp 
1985). The overlap in both size and plumage colour between the sexes 
is considerable, however, and the differences become apparent only 
when a series of specimens is studied. No sexual dimorphism has been 
described in the subspecies U. e. saturata. 

In the subspecies U. e. epops, significant differences between the 
sexes were detected in the colours of the mantle and breast (‘Table 1). 
The males are significantly paler on the mantle and breast, and more 
yellowish-red on the mantle. Also in the subspecies U. e. saturata the 
males are generally paler and more yellowish-red. This result makes it 
obvious that in an analysis of differences between the subspecies the 
sexes must be treated separately. 


Seasonal variation 

It is unknown to what degree sunlight may affect plumage colour in 
the Hoopoe; and seasonal variation in the plumage colour, if large, 
would obscure the statistical analyses. In the present samples, no 


P.G. P. Ericson 21 Bull. B.O.C. 1997 117(1) 


TABLE 1 
Descriptive statistics for the samples studied and a test of sexual dimorphism in mantle 
and breast colours in the Hoopoe subspecies Upupa e. epops and U. e. saturata. L is the 
position on the grey scale (larger values mean darker individuals). The saturation 
(“colourfulness”’) is estimated by calculating chroma as (a?+b’)’, where a is the position 
on the red-green scale and 6 on the yellow-blue scale, and the hue of the colour by 
calculating arctan (b/a) 


Males Females 
n range mean s.d. m range mean s.d. i» ddyngeds 
U. e. epops 
Mantle 
L 9 48.5-58.7 53.7 3.76 8 46.5-51.4 49.0 1.79 3.241 15 <0.01 
chroma 9 15.9-19.1 19.1 2.35 8 14.6-17.9 16.0 1.09 3.442 15 <0.01 
arctan(b/a) 9 = 1.1-1.1 a O02) 8. Tei Va (0:02. 0.147" 15 ns 
Breast 
is 9 57.6-68.1 62.7 3.87 8 56.0-63.0 59.2 2.40 2.228 15 <0.05 
chroma 9 19.4-27.7 21.7 2.84 8 17.2-24.6 20.8 2.46 0.777 15 ns 
arctan(b/a) 9 1.0-1.1 i O05 8 Ep=iNi m1 O05. 0038 e015 ns 
U. e. saturata 
Mantle 
L 10 43.2-52.7 47.3 3.00 6 43.8-47.4 46.3 1.35 0.817 14 ns 
chroma 10 12.9-17.4 14.9 1.58 6 10.8-13.8 12.8 1.16 2.799 14 <0.05 
arctan(b/a) 10° 1.1-1.1 4 -$8-02+ Gf} 1.1 0.01 0.638 14 ns 
Breast 
10 52.4-62.6 57.2 3.43 6 52.3-57.5 54.8 1.97 1.539 14 ns 
chroma 10 716.2-714 18.8 » 1.58: 6 + 15.1=20:0. 17.9. 1.78 1.154 14 ns 
arctan(b/a) 10 1.0-1.1 1.0 0.03 6 1.1-1.1 1.1 0.01 3.410 14 <0.05 


statistically significant seasonal variation, measured as the correlation 
between the collecting month and the different colour variables, was 
detected. A tendency towards paler coloration later in the year seems to 
exist in male U. e. epops, but not in females nor in any sex of U. e. 
saturata. 


Differences between subspecies U. e. epops and U. e. saturata 

Males. Highly significant differences between the subspecies epops 
and saturata were found in all variables, except the arctan calculations 
for the mantle and breast (meaning that the hues are identical). As was 
to be expected, the males of U. e. epops proved to be much paler and 
more yellowish-red than those of U. e. saturata (Table 2), and a 
bivariate plot of the chroma-variables expressing the saturation 
provides a very good separation between the samples (Fig. 1). 

Females. The females of the two subspecies also differ significantly in 
many variables. The most significant differences were found in the 
mantle, U. e. epops being on average the palest and most yellowish-red 


(Table 2). 


The Swedish Hoopoes 
The Swedish Hoopoes are very heterogeneous in regard to their 
general coloration. By adding the sex-determined Swedish individuals 


P.G. P. Ericson 22 Bull. B.O.C. 1997 117(1) 


TABLE 2 
Differences in the colour of the mantle and breast between the Hoopoe subspecies Upupa 
e. epops and U. e. saturata. Descriptive statistics and colour parameters as in Table 1. 


t d.f. i 
Males 
Mantle 
iD 4.103 1-7 <0.01 
chroma 4.627 17 <0.001 
arctan(b/a) 1.111 17 ns 
Breast 
L 3.282 17 <0.01 
chroma 2.900 17 <0.01 
arctan(b/a) 1.047 17 ns 
Females 
Mantle 
iL, 3.079 A2 <0.05 
chroma 5.241 12 <0.001 
arctan(b/a) 0.257 lg ns 
Breast 
i9 3.598 12 <0.01 
chroma 1.727, L2 <0.05 
arctan(b/a) 1.102 12 ns 


(two males and three females) to the bivariate chroma-plots it becomes 
obvious that both the subspecies U. e. epops and saturata are 
represented (Figs 1 and 2). Three unsexed Swedish birds are also dark 
enough to merit allocation to the subspecies U. e. saturata (see 
Appendix). The palest and most yellowish-red individuals (U. e. epops) 
are found in the spring while the darker (U. e. saturata) are generally 
found in the autumn. One individual (NRM 760184) that is definitely 
U. e. saturata was found in May 1975 and may well constitute the third 
known case of a Hoopoe surviving a winter in Sweden. Interestingly, 
one of the two previous records is from the same winter, 1974-75 


(Risberg 1979). 


Discussion 


Two points are fundamental to the following discussion. (1) The 
validity of U. e. saturata as a distinctive subspecies, which has 
sometimes been questioned (Kozlova 1932, Dement’ev et al. 1951) 
although recognised by most authors, can be confirmed. (2) Neither the 
material studied, nor any published information, suggests that 
individuals approaching U. e. saturata in plumage darkness occur 
anywhere within the range of U. e. epops, except for in the Ob—Yenisey 
watershed where the two forms intergrade. 

Hoopoes occurring in Scandinavia in the spring are likely to be 
mostly southeastern European birds that have prolonged their 
northward migration from the winter quarters slightly too far north 


P.G. P: Ericson 23 Bull. B.O.C. 1997 117(1) 


Breast 


Mantle 


Figure 1. Plot of the chroma-values calculated for the mantle and breast, respectively, in 
male Hoopoes. A high score along the x-axis indicates that the individual has a more 
reddish plumage, and along the y-axis that the plumage is more yellowish. Open circles 
denote Upupa epops epops, filled circles U. e. saturata, crosses Swedish male Hoopoes. 


Breast 


10 15 20 


Mantle 


Figure 2. Plot of the chroma-values calculated for the mantle and breast, respectively, in 
female Hoopoes. A high score along the x-axis indicates that the individual has a more 
reddish plumage, and along the y-axis that the plumage is more yellowish. Open circles 
denote Upupa epops epops, filled circles U. e. saturata, crosses Swedish female Hoopoes. 


P. G. P. Ericson aa Bull. B.O.C. 1997 117(1) 


Figure 3. Spring (left-hand) and autumn (right-hand) records of Hoopoes in Sweden in 
1984 (from Risberg 1985). 


(Otterlind 1954). As predicted by this theory, most Hoopoes collected 
in spring in the NRM collection are typical nominate epops. The 
autumn records of Hoopoes in Sweden show a distributional pattern 
very different from that of the spring records (Fig. 3). This observation 
led Risberg (1979) to assume that the autumn birds have a different 
geographical origin from that of the spring birds. He suggested that 
they may derive from a population breeding north of the Caspian and 
Aral Seas from which some individuals reach Sweden due to reversed 
migration. Hoopoes of this population normally winter in India but 
some birds might unintentionally migrate on a reciprocal course. What 


P.G. P. Ericson 25 Bull. B.O.C. 1997 117(1) 


Risberg did not know is that some (many?) autumn Hoopoes are 
morphologically different from nominate U. e. epops, to which the 
population north of the Caspian and Aral Seas belongs. In this part of 
its range, as in all other parts, U. e. epops is rather pale and 
yellowish-reddish (Vaurie 1965). The two individuals from this 
population in the collection of the Swedish Museum of Natural History 
(NRM 566782 and 566783) confirm this by being among the palest 
of all U. e. epops individuals studied. In their mantle colour they both 
fall clearly outside the range of U. e. saturata. ‘The occurrence of 
the subspecies U. e. saturata in Sweden can thus not be explained 
by reversed migration from the population north of the Caspian and 
Aral Seas. 

If the Swedish occurrence of U. e. saturata is to be explained by 
reversed migration a much more easterly population, well east of the 
Yenisey river, must be involved. It is possible that Hoopoes with a 
reversed migration from such a population migrate west through the 
extensive forest-steppe of southern Russia and neighbouring countries 
when confronted by the clearly unfavourable habitat of the forested 
Siberian taiga to the north. 


Acknowledgements 


The impetus to this work came from Mr. Gunnar Stré6mberg who forwarded a Hoopoe 
dead in Blekinge in November 1993 to the Swedish Museum of Natural History with a 
note concerning its unusually dark plumage. This excellent observation is here gratefully 
acknowledged. I have benefitted greatly from the knowledge and skill of Dr. Staffan 
Andersson, who taught me how to use the spectrometer and to avoid many pitfalls. 
Mr. Francisco Hernandez Carrasquilla helped me during the long measuring sessions. 
I also thank Dr. Staffan Andersson and Mr. Goran Frisk for commenting on the 
manuscript. 


References: 

CIE 1986. CIE Publication, No. 15.2. Color. 2nd edn. 

Cramp, S. (ed.) 1985. The Birds of the Western Palearctic. Vol. 4. Oxford Univ. Press. 

Dement’ev, G. P., Gladkov, N. A., Ptushenko, E. S., Spangenberg, E. P. & 
Sudilovskaya, A. M. 1951. Birds of the Soviet Union. Moskva. 

Kozlova, E. V. 1932. The birds of south-west Transbaikalia, northern Mongolia, and 
central Gobi, pt 3. Ibis (13)2: 567-596. 

Lénnberg, E. 1909. Notes on birds collected by Mr. Otto Bamberg in southern 
Transbaicalia and northern Mongolia. Arkiv fér Zoologi 5: 1-42. 

Otterlind, G. 1954. Flyttning och utbredning. Ett bidrag till kannedomen om den 
skandinaviska fagelfaunans utbredningsdynamik. Vdr Fadgelvdrld 13: 83-113. 

Pettersson, J. 1994. Harfageln hackar aterigen pa Oland. Calidris 23: 54-58. 

Risberg, L. 1979. Harfagelns Upupa epops forekomst i Sverige 1958-77. Var Fagelvdarld 
38: 221-230. 

Risberg, L. 1985. Fagelrapport f6r 1984. Var Fageludrld 44: 391-426. 

SOF 1990. Sveriges Faglar. 2nd edn. Stockholm. 

Stromberg, G. 1994. Sen harfagel i Blekinge. Var Fagelvdrld 53: 36. 

Vaurie, C. 1959. Systematic notes on Palearctic birds. No. 38 Alcedinidae, Meropidae, 
Upupidae, and Apodidae. Am. Mus. Novit., 1971. 

Vaurie, C. 1965. The Birds of the Palearctic Fauna: Non-Passeriformes. London. 


Address: Dr Per Ericson, Swedish Museum of Natural History, P.O. Box 50007, 
S-104 05 Stockholm, Sweden. 


© British Ornithologists’ Club 1997 


PGs Pt 


NRM 
no. 


566788 
566802 
566803 
690176 
690175 
566784 
690177 
566782 
566789 
566790 
566804 
566781 
566780 
566783 
566786 
566787 
566785 
566797 
566798 
566800 
556443 
556445 
556447 
566795 
566796 
566791 
566792 
566801 
566799 
556444 
556446 
566794 
566793 
760184 
786231 
760006 
906010 
786154 
740041 
763077 
936991 


Ericson 


List of Hoopoes Upupa epops examined 


Subspecies Sex 


epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
epops 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata 
saturata* 
epops* 
saturata* 
saturata* 
epops* 
saturata** 
saturata** 
saturata** 


vuve amma menemeS SSS SS ISPS rere eres SESS BESS 


26 


Appendix 


Locality 


Algeria, Quargla 

Canary, Fuertaventura Isl. 
Canary, Fuertaventura Isl., Oliva 
Hungary, Csomad 
Hungary, Dunakeszi 
Hungary, Moina Azecréod 
Hungary, Urb6 

S.E. Russia 

W. Russia, Slonim 
Algeria, Lambése 
Canary, Fuertaventura Isl., Oliva 
Romania 

Russ. Turkestan, Baimgol 
S.E. Russia 

Tunisia, Bir Mrabot 
Tunisia, El Bered 
Tunisia, Sidi Mansour 
China, W. Shansi Prov. 
China, W. Shansi Prov. 
China, W. Shansi Prov. 
Korea, Riuganpo 

Korea, Shuotsu 

Korea, Shuotsu 
Mongolia, Tabool 
Mongolia, Tabool 

S. Transbaicalia, Kjachta 
S. Transbaicalia, Kjachta 
China, Richthofen Mts. 
China, W. Shansi Prov. 
Korea, Riuganpo 

Korea, Shuotsu 
Mongolia, Tabool 

S. Transbaicalia, Kjachta 


Sweden, Vastmanland, Lindesberg 


Sweden, Varmland, Svaneholm 
Sweden, Skane, Helsingborg 
Sweden, Dalarna, Rattvik 
Sweden, Sédermanland, Dunker 
Sweden, Smaland, Vastervik 
Sweden, Smaland, Vetlanda 
Sweden, Blekinge, Sturk6 


Coll. date 


5 Sep. 1912 
Mar. 1912 
Feb. 1910 

4 Jul. 1965 

10 Aug. 1965 


29 Aug. 1965 
4 Mar. 1909 
12 Jul. 1916 
7 May 1910 
Feb. 1910 
20 May 1906 
10 May 1902 
4 May 1909 
28 Mar. 1906 
4 Mar. 1909 
28 Mar. 1903 
24 May 1921 
16 May 1921 
27 May 1921 
13 May 1936 
1 Jul. 1935 
5 Jul. 1935 
30 Jul. 1919 
30 Jul. 1919 
15 May 1908 
20 May 1908 
2 May 1932 
16 Apr. 1921 
23 Apr. 1936 
4 Jul. 1935 
21 Jul. 1919 
13 May 1908 
12 May 1975 
18 May 1978 
4 Jan. 1976 
16 Dec. 1990 
23 Apr. 1978 
20 Oct. 1973 
25 Jan. 1976 
25 Nov. 1993 


Bull. B.O.C. 1997 117(1) 


Remarks 


Type 
Paratype 


Paratype 


* Allocation to this subspecies based on the spectrometry measurements in this study 
**not measured by the spectrometer but tentatively allocated to this subspecies 


M. Galetti et al. 27 Bull. B.O.C. 1997 117(1) 


Records of Harpy and Crested Eagles in the 
Brazilian Atlantic forest 


by Mauro Galetti, Paulo Martuscelli, Marco Aurélio Pizo 
&F Isaac Simao 


Received 16 March 1996 


Forest raptor populations are declining throughout the world, and 
information on the ecology and even the distribution of most species is 
still lacking (Bierregaard 1995; see also del Hoyo et al. 1994). The main 
threats for raptors are deforestation, hunting and competition with 
humans for prey (Thiollay 1984, Redford 1992). The Atlantic forest of 
Brazil once covered 1 million km’, of which only about 5% now 
remains; it holds a vast diversity of endemic birds and mammals and is 
considered one of the most endangered ecosystems of the world (SOS 
Mata Atlantica & INPE 1992). 

The Harpy Eagle Harpia harpyja is the largest raptor in America and 
one of the largest in the world. It occurs from México to Bolivia 
and Argentina, and in a large part of Brazil (Sick 1993). Its ecology and 
distribution is very poorly known and most studies have been located 
in the Amazon region (Rettig 1978, Izor 1985). Harpy Eagles are 
considered scarce in the Atlantic forest and recent records are few 
(Chebez et al. 1990, Albuquerque 1995). 

The Crested Eagle Morphnus guianensis is similar to an immature 
Harpy Eagle but the crest is not divided into two ‘horns’ (Sick 1993). It 
occurs from Guatemala to Bolivia and Argentina (Misiones), and there 
are records from most of Brazil (Sick 1993). Very little is known about 
its ecology and distribution (Bierregaard 1984). 

This paper presents six recent records of Harpy Eagle and four of 
Crested Eagle in the Atlantic forest of Brazil, and suggests a possible 
migratory population of Harpy Eagles in the south of its range. 


Observations 


Harpy Eagle 

On 25 July 1991 a Harpy Eagle was observed around 15.00h at 
Estagao Experimental Pau-Brasil, Porto Seguro, south of Bahia 
(16°19’S, 39°11’W). The area is a reserve of 1145 ha covered mainly by 
mature forest and secondary vegetation (Mori et al. 1983). The eagle 
was a mature individual perched in a dead tree in logged secondary 
vegetation. Possible prey for large eagles that occur in this area are the 
monkeys Callithrix geoffroyi and Cebus apella, and the sloth Bradypus 
variegatus (Oliver & Santos 1991). 

On 11 August 1992, an adult Harpy Eagle was observed just after it 
killed a sloth (Bradypus variegatus) in the Compania Vale do Rio Doce 
Reserve (CVRD), Linhares, Espirito Santo (19°06’'S, 39°45’W). The 
eagle was perched only 5 m up and flew away on being observed. On 
18 December 1992 probably the same individual was observed, perched 


M. Galetti et al. 28 Bull. B.O.C. 1997 117(4) 


TABLE 1 
Comparative density, expressed as no. of groups/km/7, of arboreal prey of large raptors in 
two localities of the Atlantic forest 


Area Forested Density 

(State) area (km*) Bradypus Cebus Alouatta Callicebus Callithrix Potos 
Linhares* (E.S.') 157 ? 2.19 + angi |) 10.46 ? 
Intervales** (S.P.7) 490 — 0.16 aa — = — 


*data from Chiarello (1995); **data from M. Galetti in prep. 

'Total forested area in the state is 4,023 km. 

Total forested area in the state is ith, 314 km? 

Abundance: +=occurs but extremely rare; —-=does not occur; ?=occurs but density 
unknown. 


at a height of 20m. In 1995 a Harpy Eagle nest was found in a 
lecythidaceous tree in CVRD (R. Jesus pers. comm.). Faunal surveys 
in other areas of Espirito Santo state have not recorded Harpy Eagles 
(A. G. Chiarello pers. comm.). The CVRD reserve holds a high density 
and diversity of prey for large predators (both mammalian and avian), 
including agoutis, pacas and monkeys, as well as large birds (Chiarello 
1995, Scott & Brooke 1985; see Table 1). 

On 29 January 1990 a claw of a Harpy Eagle was confiscated from a 
hunter. The hunter stated that the eagle was killed in July 1989 at 
Varadouro, Cananéia, S40 Paulo (25°15’S, 48°05’W). On 1 July 1990 
two adult Harpy Eagles were observed soaring at Ariri, on the south 
coast of Sao Paulo state (25°03’S, 48°07’W). In July 1991 and 1993 two 
Harpy Eagles were observed further north in the Cananéia region 
within 45 km” of continuous forest. The eagles were observed there 
during the entire month. This region is covered by pristine Atlantic 
forest, mangroves and restinga, ranging from sea level to 1350m 
altitude. 


Crested Eagle 

On 25 May 1990 an adult Crested Eagle was observed soaring over 
Parque Estadual Jacupiranga at 750 m a.s.1. (24°53’S, 48°22'W). Also in 
the same park but at 50 maz.s.]. an eagle was observed on 14 December 
1992. This park covers 1100km* of Atlantic forest ranging from 
lowland to montane forest. 

On 14 December 1992 another adult Crested Eagle was observed in 
Parque Estadual do Morro do Diabo (see Valladares-Padua & Cullen 
1994 for site description), in semideciduous forest close to the Rio 
Paranapanema. This area contains 320km* of semideciduous and 
riverine forest and is the largest fragment of forest remaining in the 
hinterland of Sao Paulo. 

At 17.05h on 24 February 1995, a Crested Eagle was observed 
perched 20m up at Carmo station (700m a.s.l.), Parque Estadual 
Intervales (24°14’S, 48°04’W), near Capao Bonito, Sao Paulo. This 
reserve covers 490 km? of secondary and primary Atlantic forest, which 


M. Galetti et al. 29 Bull. B.OLC3199%.117(4) 


is continuous with the Parque Estadual Carlos Botelho and Petar (see 
Pizo et al. 1995). Large prey species (monkeys, agoutis, deer) for large 
predators are very scarce in this area (M. Galetti unpubl. data; ‘Table 1). 


Discussion 


Both Harpy and Crested Eagles need large areas for survival. Thiollay 
(1989) estimated that a pair of either species may have a home range 
more than 100km*. The Harpy Eagle’s diet usually includes game 
species, such as monkeys (Alouatta, Cebus), sloths (Bradypus), deer 
(Mazama) and also large birds (cracids, macaws, seriemas) (Rettig 
1978, Izor 1985, Sick 1993). Crested Eagles prey mainly on snakes, 
large birds (trumpeters, cock-of-the-rock) and occasionally arboreal 
mammals such as kinkajous and young spider monkeys (Bierregaard 
1984, Julliot 1994; see also del Hoyo et al. 1994). 

The remaining, highly fragmented Atlantic forest covers c. 
60 000 km? (SOS Mata Atlantica & INPE 1992). The size of this area 
suggests there is still suitable habitat for large raptors, but the area per 
se cannot predict the abundance of large raptors in this forest. Thiollay 
(1984) suggests that the human impact of hunting on game animals can 
reduce the prey availability for large raptors, and the Harpy Eagle 
would be the first species to disappear in hunted forests. In fact, 
hunting pressure in Neotropical forests can extirpate most game species 
in a few years (Peres 1990). 

The remaining forested area in Sao Paulo (c. 17 314 km’) is much 
greater than in Espirito Santo (c. 4023 km’, data from SOS Mata 
Atlantica & INPE 1992). Harpy Eagles have been recorded recently in 
Sao Paulo only during the winter (June, July and August), suggesting 
migratory movements (see Sick 1993). Migratory movement is also 
suggested by the date of collection of earlier specimens of Harpy Eagles 
in Sao Paulo (MZUSP, two eagles collected in August, one in Ituverava 
and another in Juquia). Albuquerque (1995) also recorded the Harpy 
Eagle only in the winter (or after the passage of a cold air mass) in Serra 
do Tabuleiro, Santa Catarina. Straube (1989) suggests that the Atlantic 
avifauna colonized Misiones via riverine forest of the Iguacu river. 
Harpy Eagles might be expected to migrate along the forests fringing 
the Paranapanema river. 

Harpy Eagles are known to breed in Misiones, Argentina (Chebez 
et al. 1990), and have been recorded there as recently as January 1996 
(E. R. Krauczuck im litt.). Three nests in Argentina were located in 
sites protected from cold winds from the south, and the eagles probably 
migrate during cold winters (E. R. Krauczuck in litt.). Chebez (1994) 
suggests that Parque Provincial Urugua-i and Yaboti are the only 
places that still have Harpy Eagles in Argentina. 

The rarity of Harpy Eagles in the Atlantic forests of SAo Paulo (and 
probably Santa Catarina and Parana) can be explained not only by 
habitat loss but also by the density of their prey. Large mammal prey 
(mainly primates and sloths) have low densities in most large reserves 
in southeastern Brazil (Chiarello & Galetti 1994, M. Galetti unpubl. 
data; Table 1). The exceptions are the north of Espirito Santo and 


M. Galetti et al. 30 Bull. B.O.C. 1997 117(1) 


south of Bahia where the density of monkeys, agoutis and other 
possible prey of Harpy Eagles is high (Oliver & Santos 1991, Chiarello 
1995, A. Chiarello unpubl. data). It is not known why the density of 
arboreal mammals in Espirito Santo and Bahia is so different from S4o 
Paulo; both areas are affected heavily by hunting. In CVRD, Espirito 
Santo, Harpy Eagles have been observed since the 1970s, and it is one 
of the few places in the whole Brazilian Atlantic forest where this eagle 
is known to breed. Houston (1987) found a high correlation between 
the density of mammals and vultures in Neotropical forests, and the 
same is probably true for large raptors as well (see also Thiollay 1984). 

The data presented here suggest that there are at least two 
populations of Harpy Eagles in the Atlantic forest: sedentary 
individuals in the north of Espirito Santo and south of Bahia, and 
migratory ones in the south of Brazil (Sdo Paulo, Parana and Santa 
Catarina), breeding in Misiones, Argentina. 

The Crested Eagle is considered naturally rare (Sick 1993), and the 
only previous record of this species for Sao Paulo was in 1900 (MZUSP 
2417 from Apiai). There has been a recent report from Barrolandia, 
near Porto Seguro Bahia (Anon. 1995). It is difficult to know why the 
Crested Eagle is so rare in southeastern Brazil, but arboreal snakes 
(boas), their main prey, are extremely rare in the Atlantic forest of Sao 
Paulo (pers. obs.). 

‘To sum up, the status of Harpy and Crested Eagles in the Atlantic 
forest is highly endangered. Large raptors need more than large areas to 
survive; it is also necessary to maintain high levels of their prey (see 
Redford 1992). Most reserves in the Brazilian Atlantic forests are 
‘empty forests’ where hunting is still a common practice (Oliver & 
Santos 1991, Galetti & Chivers 1995). Several reserves still have a high 
human population, hunting and logging in the forest, and the 
boundaries of many reserves are just virtual. 

The conservation of Harpy Eagles should be given special attention 
if migratory movements are confirmed (probably possible by radio- 
telemetry only). Also it is paramount to protect the forests in northern 
Espirito Santo and southern Bahia where Harpy Eagles are resident. 
Oliver & Santos (1991) suggest several areas where new reserves could 
be established. Harpy and Crested Eagles may also occur in other 
remaining large forest blocks (e.g. Parque Nacional de Monte Pascoal), 
and these should be investigated. 


Acknowledgements 


We are grateful to Fundaciao Florestal and Instituto Florestal for permission to visit their 
reserves in Sao Paulo, to CVRD for logistical support at Linhares, to FMB for constant 
support to our ornithological projects, to F. Olmos, K. McConkey and A. D. Cuaron for 
critical review of the manuscript, and E. R. Krauczuk for his unpublished data about the 
status of Harpy Eagles in Argentina. M. Galetti is supported by CNPq, Marco A. Pizo by 
CAPES and FAPESP, I. Simao by FAPESP and CAPES. 


References: 

Albuquerque, J. L. B. 1995. Observations of rare raptors in southern Atlantic rain-forest 
of Brazil. 7. Field Orn. 66: 363-369. 

Anonymous. 1995. Neotropical notebook. Cotinga 4: 68. 


M. Galetti et al. 31 Bull. B.O.C. 1997 117(1) 


Bierregaard, R. O. 1984. Observations of the nesting biology of the Guiana Crested Eagle 
(Morphnus guianensis). Wilson Bull. 96: 1-5. 

Bierregaard, R. O. 1995. The biology and conservation status of Central and South 
American Falconiformes: a survey of current knowledge. Bird Cons. Int. 5: 325-340. 

Chebez, J. C. 1994. Harpia. Los que se van. Edit. Albatroz. 

Chebez, J. C., Crome, M. S., Serret, A. & Taborda, A. 1990. La nidificacion de la Harpia 
(Harpia harpyja) en Argentina. El Hornero 13: 155-158. 

Chiarello, A. G. 1995. Density and habitat use of primates at an Atlantic forest reserve of 
southeastern Brazil. Rev. Brasil. Biol. 55: 105-110. 

Chiarello, A. G. & Galetti, M. 1994. Conservation of the brown howler monkey in 
southeastern Brazil. Oryx 28: 37-42. 

del Hoyo, J., Elliot, A. & Sargatal, J. 1994. Handbook of the Birds of the World. Vol. 2. 
Lynx Edicions, Barcelona. 

Galetti, M. & Chivers, D. J. 1995. Palm harvest threatens Brazil’s best protected area of 
Atlantic forest. Oryx 29: 225-226. 

Houston, D. C. 1987. The effect of reduced mammal numbers on Cathartes vultures in 
Neotropical forests. Biol. Conserv. 41: 91-98. 

Izor, R. J. 1985. Sloths and other mammalian prey of the Harpy Eagle. In: G. G. 
Montgomery (ed.) The Conservation and Ecology of Armadillos, Sloths, and 
Vermilinguas. Smithsonian Institution Press, Washington. 

Julliot, C. 1994. Predation of a young spider monkey (Ateles paniscus) by a crested eagle 
(Morphnus guianensis). Folia primatol. 63: 75-77. 

Mori, S. A., Boom, B. M., Santo, A. M. & Santos, T. 1983. Southern Bahian moist 
forests. Bot. Review 49: 155-232. 

Oliver, W. L. R. & Santos, I. B. 1991. Threatened endemic mammals of the Atlantic 
forest region of south-east Brazil. Wildlife Preservation Trust. Special Scientific 
Report no 4. 

Peres, C. A. 1990. Effects of hunting on western Amazonian primate communities. Biol. 
Conserv. 54: 47-59. 

Pizo, M. A., Simao, I. & Galetti, M. 1995. Diet and flock size of sympatric parrots in the 
Atlantic forest of Brazil. Ornitologia Neotropical 6: 87-95. 

Redford, K. H. 1992. The empty forest. Bioscience 42: 412-422. 

Rettig, N. L. 1978. Breeding behavior of the Harpy Eagle (Harpia harpyja). Auk 95: 
628-643. 

Scott, D. A. & Brooke, M de L. 1985. The endangered avifauna of southeastern Brazil: a 
report on the BOU/WWF Expedition of 1980/81 and 1981/82. Pp. 115-139 in 
Conservation of Tropical Birds. 1CBP Technical Publication no 4. 

Sick, H. 1993. Birds in Brazil. Princeton Univ. Press. 

SOS Mata Atlantica & INPE. 1992. Atlas da evolugado dos remanescentes florestais e 
ecosistemas associados do dominio da Mata Atlantica no periodo de 1985-1990. 
Fundacgao SOS Mata Atlantica, Sao Paulo. 

Straube, F. C. 1989. Sobre a distribuigéo do Macropsalis creagra no estado do Parana. 
Sulornis 10: 12-21. 

Thiollay, J. M. 1981. Raptor community structure of a primary rain forest in French 
Guiana and effect of human hunting pressure. Raptor Res. 18: 117-122. 

Thiollay, J. M. 1989. Area requirements for the conservation of rain forest raptors and 
game birds in French Guiana. Conserv. Biol. 3: 128-137. 

Valladares-Padua, C. & Cullen, L. 1994. Distribution, abundance and minimum viable 


metapopulation of the black lion tamarin Leonthopithecus chrysopygus. Dodo 30: 
80-88. 


Address: Mauro Galetti, Wildlife Research Group, Department of Anatomy, University 
of Cambridge, Cambridge CB2 3DY, U.K. Paulo Martuscelli, C.P. 194, 11750-970 
Peruibe, S.P., Brazil. Marco A. Pizo and Isaac Simao, Departamento de Zoologia, 
UNICAMP, C.P. 6109, 13081-970 Campinas, Sao Paulo, Brazil. 


© British Ornithologists’ Club 1997 


T. Brooks & G. Dutson S2 Bull. B.O.C. 1997 117(1) 


‘Twenty-nine new island records of birds from 
the Philippines 


by Thomas Brooks & Guy Dutson 
Received 19 November 1995 


We spent 21 July to 19 August 1994 birdwatching on the Philippine 
islands of Bohol, Cebu, Panay, Sanga-Sanga, Bongao, Tawi-Tawi, 
Mindanao and Mindoro. During this period, we obtained sight records 
of a number of bird species not previously known from these islands, 
according to the distributional information given in Dickinson et al. 
(1991). Field-notes supporting the identification of these birds are 
available from the authors. 


LITTLE GREBE Tachybaptus ruficollis 

‘Twelve birds, including several immatures, were seen on a small lake 
just east of Sanga-Sanga town, Sanga-Sanga, on 10 August. The 
species has not previously been recorded in the Sulu Archipelago, 


although it is known from Mindanao, and from Borneo (MacKinnon & 
Phillipps 1993). 


WHISTLING-DUCK SP. Dendrocygna sp. 

A single bird, almost certainly Wandering Whistling-Duck Dendro- 
cygna arcuata, was seen in flight at dusk in Barangay Magsagaw, 
Tawi-Tawi, on 8 August. The species is common on Sanga-Sanga, but 
is otherwise known from elsewhere in the Sulu Archipelago only from 
the sight records of duPont & Rabor (1973). The possible confusion 
species, Spotted Whistling-Duck D. guttata, is known from the 
south-west Philippines (Mindanao, Basilan and Jolo) from a few 
historical specimens. 


HAWKE SP. Accipiter sp. 

A small Accipiter sp. was flushed from secondary forest in Barangay 
Alojipan, at the base of Mt Madja-as, Panay, on 2 August. Our views 
were insufficient to identify the bird, but no Accipiter spp. are 
previously known from Panay. 


PEREGRINE FALCON Falco peregrinus 

One bird, probably a female, of the dark resident race F. p. ernesti 
was watched fiercely mobbing a White-bellied Sea-Eagle Haliaeetus 
leucogaster on Bongao peak on 6 August. The species is not listed for 
the Sulu Archipelago by Dickinson et al. (1991), although it was 
recorded on Tawi-Tawi on 23 March 1987 by Hornskov (1995). Birds 
are known to be resident on Mindanao and Cagayan Sulu, and in 
northern Borneo (MacKinnon & Phillipps 1993). 


TABON SCRUBFOWL Megapodius cumingit 


A single bird was seen in dense forest on Bongao peak at dawn on 
6 August. Although not listed for the island by Dickinson et al. (1991), 


T. Brooks & G. Dutson 33 Bull. B.O.C. 1997 117(1) 


the species was seen here by I. Gardner (27m /itt. 1993). It is known from 
most islands in the Sulu Archipelago. 


BUTTONQUAIL SP. Turnix sp. 

One Turnix sp. was flushed from grass on Sanga-Sanga on 
10 August, but the views obtained were insufficient for specific 
identification. The only Turnix sp. known from the Sulu Archipelago 
is Small Buttonquail JT. sylvatica, which is itself only known from 
the type of T. s. suluensis, collected on Jolo in 1903 (Mearns 1905). 
T. sylvatica is also the only Turnix sp. known from Mindanao, and no 
species occurs on Borneo (MacKinnon & Phillipps 1993). 


RUDDY-BREASTED CRAKE Porzana fusca 

A single bird was seen in dense secondary riverine forest 
undergrowth at c. 100m altitude in the Sampulong Bolo Natural 
Sanctuary, Sara, Panay, on 31 July. The species is known from most of 
the large Philippine islands. 


COMMON SANDPIPER Actitis hypoleucos 

Three birds were seen flying along the beach in Sanga-Sanga town, 
Sanga-Sanga, on 5 August. The species is a common passage migrant 
throughout the Philippines. It is not listed for Sanga-Sanga by 
Dickinson et al. (1991) although six were recorded here in October 
1971 by duPont & Rabor (1973) and three on 24 March 1987 by 
S. Jensen and J. Hornskov (zn /itt. 1992). 


WOOD SANDPIPER 77inga glareola 


One bird was seen on the northern shore of Bongao on 13 August. 
‘The species is a common passge migrant to most Philippine islands. 


ORIENTAL PRATINCOLE Glareola maldivarum 


A single bird was seen hawking over paddyfields near Concepcion, 
Mindoro, on 18 August. Although not listed in Dickinson et al. (1991), 
I. Gardner (in litt. 1988) recorded a single bird near San José in March 
1988 and Evans et al. (1993) recorded two birds near Malpalon on 
24 September 1991. The species has not previously been recorded in 
the Philippines in August. 


COMMON TERN Sterna hirundo 


Two birds were seen just off the east coast of Tawi-Tawi on 
12 August. The species is an uncommon passage migrant throughout 
the Philippines, and although not listed for Tawi-Tawi by Dickinson 
et al. (1991), about 50 were recorded off the east coast on 28 March 
1987 by S. Jensen and J. Hornskov (in litt. 1992). 


BLACK-NAPED TERN Sterna sumatrana 

Three adult birds were seen just offshore from Cebu City, Cebu, 
from the Tagbilaran to Cebu ferry, on 27 July. The species is likely to 
be an uncommon resident throughout the Philippines, although 


T. Brooks & G. Dutson 34 Bull. BOC 1997 Tiait) 


Hornskov (1995) suggests that a recent range extension may have taken 
place. 


SPOTTED DOVE Streptopelia chinensis 
This species was commonly seen in agricultural land at and near the 
Sablayan Penal Colony, Mindoro, 15-18 August. Although not listed 


for the island in Dickinson et al. (1991) it was commonly recorded in 
1988 by I. Gardner (zn litt. 1988) and in 1991 by Evans ez al. (1993). 


ZEBRA DOVE Geopelia striata | 

The distinctive calls of this species were heard on Sanga-Sanga on 
12 August. Escaped cagebirds are known to have colonised a number of 
Philippine islands (Dickinson et al. 1991, Hornskov 1995), and this 
may be the case on Sanga-Sanga. Four individuals were also seen on 
Bongao on 5 August. 


COMMON KOEL Eudynamys scolopacea 

Three individuals were seen and heard on Sanga-Sanga on 12 
August. The species is already known from many of the islands of the 
Sulu Archipelago, and was reported from Sanga-Sanga by duPont & 
Rabor (1973). 


PHILIPPINE EAGLE-OWL Bubo philippensis 

One bird was flushed at c. 15m range in deep forest at midday on 
26 July, in Rajah Sikatuna National Park, Bohol. It was a huge owl, 
about twice the bulk of a Tawny Owl Strix aluco with similar 
short-winged proportions. The plumage was warm brown with 
prominently barred tail and remiges, paler primary bases and a dark 
carpal patch. This is the only large owl in the Philippines other than the 
Spotted Wood-Owl Strix seloputo of Palawan, which is considerably 
smaller and has large pale spots on its upperparts. A large owl probably 
of this species was also seen in Rajah Sikatuna National Park on 
28 January 1994 (J. Hornbuckle & P. Morris in litt.). The species is 
endemic to the eastern Philippines (known from Luzon, Catanduanes, 
Samar, Leyte and Mindanao) and is thus not unexpected on Bohol. 


GREY-RUMPED TREESWIFT Hemuiprocne longipennis 

G.D., who is familiar with the species, saw two birds on 10 August 
and three (which were photographed) at close range on 11 August, all in 
Barangay Buan, Tawi-Tawi. This is only the second Philippine record, 
the other being of two collected on Sibutu on 2 November 1971 
(duPont & Rabor 1973). The species is common on_ Borneo 
(MacKinnon & Phillipps 1993), and its range seems to be expanding, 
for it has also recently been recorded for the first time on Lombok, 
Indonesia (Johnstone et al. 1993). 


SWIFTLET SP. Collocalia sp. 

A number of large, dark swiftlets seen between 700 and 1000 m on 
Mt Madja-as, Panay, on 2 August seem likely to have been Philippine 
Swiftlets Collocalia mearnsi. Although the species is dubiously 


T. Brooks & G. Dutson 35 Bull. B.O.C. 1997 117(1) 


separable from Island Swiftlet C. [vanikorensis] amelis in the field, 
C. mearnsi occurs on the adjacent island of Negros (C. [vanikorensts ] 
amelis is apparently absent from both Negros and Panay), and is largely 
an upland species while C. [vantkorensis] amelis is generally confined to 
the lowlands. 


SWIFTLET SP. Collocalia sp. 

Small, dark swiftlets were commonly seen on Sanga-Sanga (5, 6, 
10-13 August), with a maximum of about 50 individuals. They seem 
likely to have been Edible-nest Swiftlet Collocalia fuciphaga, which was 
also reported from the Sulu Archipelago by S. Jensen and J. Hornskov 
(in litt. 1992), but note that this record, of ten birds near Sangka- 
Sangka bridge on 24 March 1987, was subsequently accidentally listed 
as being of German’s Swiftlet Collocalia germani in Hornskov (1995). 
The species has not previously been recorded in the Sulus, although it 
is common in Borneo (MacKinnon & Phillipps 1993). 

A few birds (maximum ten individuals) were also seen on Bongao on 
5 and 6 August. The bamboo ladders on the north face of Bongao peak 
suggest that edible swiftlet nests are harvested here, and so the birds 
were probably Edible-nest Swiftlets Collocalia fuciphaga. Medway 
(1966) also received reports of the collection of edible swiftlet nests in 
the Sulus, and so although Dickinson et al. (1991) suggested that these 
reports must have originated from Cagayan Sulu, they may after all 
refer to the Sulu Archipelago proper. 


GLOSSY SWIFTLET Collocalia esculenta 

Birds were seen commonly on Sanga-Sanga (5, 6, 10-13 August). 
The species is known from Bongao but not from elsewhere in the Sulu 
Archipelago.The species was also very common on Tawi-Tawi with 
flocks of over 100 individuals seen on 6-12 August. Although not listed 
for the island by Dickinson et al. (1991), it was recorded here by 
S. Jensen and J. Hornskov (in litt. 1992) on 25 and 26 March 1987. 


PHILIPPINE NEEDLETAIL Mearnsia picina 
A single bird was seen on Sanga-Sanga on 12 August. The species 


was only recently recorded in the Sulu Archipelago for the first time, 
on Tawi-Tawi (Lambert 1993). 


HOUSE SWIFT Apus affinis 

A single bird was seen on Sanga-Sanga on 12 August. Good views 
were obtained, excluding the possibility that this was a Fork-tailed 
Swift A. pacificus. Although only known from a few islands in the 
northern Philippines, A. affinis is common on Borneo (MacKinnon & 
Phillipps 1993), and seems to be expanding in the Philippines (Evans 
et al. 1993). 


PURPLE NEEDLETAIL Hirundapus celebensis 

Four individuals were seen at 600m on Mt Madja-as, Panay, on 
2 August. The species is known from most of the larger Philippine 
islands. 


T. Brooks & G. Dutson 36 Bull. B.O.C. 1997 117(1) 


BARN SWALLOW Hiirundo rustica 

‘Two birds were seen on Bongao on 5 August. The species is a 
common passage migrant and winter visitor to the Philippines. The 
earliest Philippine record listed in Dickinson et al. (1991) is 15 August. 
Evans et al. (1993) recorded two birds on Negros on 29 July 1991. 


YELLOW-VENTED BULBUL Pycnonotus gotavier 

‘Two individuals were seen in scrub on Sanga-Sanga on 10 August. 
The species is already known from most of the islands in the Sulu 
Archipelago. A single bird was also seen on Bongao on 5 August. The 
subspecies of these birds was not determined but would indicate 
whether these populations originated from Borneo or the Philippines. 


CELESTIAL MONARCH Hypothymis coelestis 

A bird seen for some minutes at close range in logged forest in 
Barangay Buan, Tawi-Tawi, on 10 and 11 August appeared to be a 
Celestial Monarch AH. coelestis but, given the range extension, may 
prove to be a new subspecies. It differed obviously from nearby 
Black-naped Monarchs H. azgurea in its pale sky-blue crown, drooping 
crest and broad eye-ring. The length of the crest was difficult to assess 
but when held away from the neck, its projection was longer than that 
of the bill. The rest of the head, nape and breast had a purple wash 
whilst the mantle and wing-coverts had sky-blue streaking. The belly 
was white and the remiges and rectrices dull blue. The similar 
Short-crested Monarch H. helenae has a shorter crest, no eye-ring, no 
purple tones and no pale streaking on the upperparts. This is the first 
record of this rare bird from the Sulu Archipelago, although it is known 
from Basilan. 


STRIPE-BREASTED RHABDORNIS Rhabdornis inornatus 

‘Two birds were seen at 900 m on Mt Madja-as, Panay, on 2 August. 
R. inornatus is known from the mountains of Negros, and Panay birds 
are presumably of the same race (R. 12. raborz). 


TREE SPARROW Passer montanus 

The species was common in Sanga-Sanga town, Sanga-Sanga (5, 6, 
10-13 August). Tree Sparrows were also common in Bongao town (5 
and 6 August), and Batu-Batu town, Tawi-Tawi (6-12 August). It has 
not been previously recorded in the Sulu Archipelago. 


SCALY-BREASTED MUNIA Lonchura punctulata 

Flocks of this species were seen in Pasonanca, Zamboanga, 
Mindanao, on 10 and 12 August. It is previously known only from the 
northern Philippines and these flocks may have originated from escaped 
cagebirds. 


Discussion 


The new island records of Philippine Eagle-Owl and of Celestial 
Monarch are most exciting and show that significant discoveries may 


T. Brooks & G. Dutson oF Bull SE .OFCe ya THe 


still be made on Philippine islands rarely visited by ornithologists. Both 
of these species are listed as “‘threatened’”’ in Collar et al. (1994). 

However, the majority of the new island records listed above are a 
depressing testimony to the rapidly continuing destruction of forest 
habitats throughout the Philippine Archipelago. We found most of 
these species to be common, and they would surely have been recorded 
during any previous ornithological work on the islands in question had 
they been present. Species such as Spotted Dove, Zebra Dove, House 
Swift, Yellow-vented Bulbul, Tree Sparrow and Scaly-breasted Munia 
are rapidly spreading throughout the country following deforestation 
and urbanization. They are replacing the endemic forest birds which 
make the avifauna of the Philippines so special, and are indicative of the 
task facing the conservation of Philippine biodiversity. 


Acknowledgements 


Our sincere thanks go to: Mr Tim Fisher and his family in Manila; the DENR staff of the 
Logarita Forest Station on Bohol; Ms Perla Magsalay and the staff of the Philippine 
Wetland and Wildlife Conservation Foundation Inc. on Cebu; Mrs Mila Ebreo and her 
family, and the staff of the DENR-Culasi on Panay; Professor Ernesto Baird and his 
family, and especially Mr Samuel Concepcion and Mr Bibie Palomo on Sanga-Sanga; the 
family of the Mayor, Mr Sadikul Sahali, and the Vice-mayor, Mr Sahabad Jaji, in 
Batu-Batu; the Barangay Captain, Mr Bidin A. Galib, and the people of Barangay 
Magsagaw who guarded us in the forest on Tawi-Tawi; the staff of the Sablayan Penal 
Colony; and Mr Leonardo Gabutero, the Chairman, and the volunteers of the Kalikasan 
Mindoro Foundation Inc., especially Mr Efrain Tejada. We also thank Mr Jon 
Hornbuckle, Mr Jesper Hornskov, Mr Pete Morris and Mr Ian Gardner for providing 
recent records, and Mr Edward Dickinson, Dr David Snow and an anonymous reviewer 
for commenting on the manuscript. TB is grateful to Dr Stuart Pimm for providing a 
travel grant. 


References: 

Collar, N. J., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to Watch 2. BirdLife 
Conservation Series No. 4. BirdLife International, Cambridge, U.K. 

Dickinson, E. C., Kennedy, R. S. & Parkes, K. C. 1991. The Birds of the Philippines. 
B.O.U. Checklist no. 12. 

duPont, J. E. & Rabor, D. S. 1973. South Sulu Archipelago birds: an expedition report. 
Nemouria 9: 1-63. 

Evans, T. D., Dutson, G. C. L. & Brooks, T. M. 1993. Cambridge Philippines Rainforest 
Project 1991. Final Report. BirdLife International Study Report no. 54. BirdLife 
International, Cambridge, U.K. 

Hornskovy, J. 1995. Recent observations of birds in the Philippine Archipelago. Forktail 
11: 1-10. 

Johnstone, R. E., van Balen, S. & Dekker, R. W. R. J. 1993. New bird records for the 
island of Lombok. Kukila 6: 124-127. 

Lambert, F. R. 1993. Some key sites and significant records of birds in the Philippines 
and Sabah. Bird Conservation International 3: 281-297. 

MacKinnon, J. & Phillipps, K. 1993. Birds of Borneo, Sumatra, Java and Bali. Oxford 
Univ. Press. 

Mearns, E. A. 1905. Descriptions of eight new Philippine birds, with notes on other 
species new to the islands. Proc. Biol. Soc. Washington 18: 83-90. 

Medway, Lord. 1966. Field characters as a guide to the specific relations of swiftlets. 
Proc. Linn. Soc. London 177: 151-172. 


Addresses: Thomas Brooks, Department of Ecology and Evolutionary Biology, 569 
Dabney Hall, University of Tennessee, Knoxville, Tennessee 37996-1610, U.S.A. 
Guy Dutson, 1 High Way, Broadstone, Dorset BH18 9NB, U.K. 


© British Ornithologists’ Club 1997 


C. B. Frith & D. W. Frith 38 Bull. B.O.C. 1997 117(1) 


‘The taxonomic status of the bird of paradise 
Paradigalla carunculata intermedia 
(Paradisaeidae) with notes on the other 
Paradigalla taxa 


by Clifford B. Frith G Dawn W. Frith 
Received 10 December 1995 


By convention, the bird of paradise genus Paradigalla (Paradisaeidae) 
comprises two species, the Long-tailed Paradigalla P. carunculata and 
the Short-tailed Paradigalla P. brevicauda (Mayr 1962, Gilliard 1969, 
Cooper & Forshaw 1977). They are medium-sized (38 and 23 cm long 
respectively), sexually monomorphic members of the subfamily 
Paradisaeinae (typical birds of paradise). Adults are jet-black over the 
entire body with strongly iridescent blue-green scale-like crown 
feathering and a subtle deep green sheen to the leading edge of 
primaries and to dense velvet-like mantle feathering. The face is 
conspicuously adorned with a yellow wattle that covers the lores and 
forehead and a bulbous blue wattle at the base of the lower mandible. 
These wattles are fully developed in P. brevicauda hatchlings, but both 
are dirty cream-yellow with a smudged greyish lower border to the 
lower mandible wattle. In much longer and acuminately-tailed adult 
P. carunculata there is an additional small bare area directly beneath 
the blue wattle that is pigmented deep orange-red; these characters, 
including the long tail, distinguish this species from smaller, truncately- 
tailed, P. brevicauda. For more detailed plumage descriptions see 
Gilliard (1969), Cooper & Forshaw (1977) and Beehler et al. (1986). 

The paradigallas occur in montane forests in New Guinea between 
approximately 1400 and 2100m asl. They are little known; an 
exception is a study of the nesting biology of P. brevicauda showing that 
only a single, presumed female, parent attended the nest (Frith & Frith 
1992). This finding suggests that males are promiscuous as in most bird 
of paradise species. 

The long-tailed P. carunculata was described by Lesson (1835) from 
a specimen of unknown location. All subsequent specimens of known 
origin are from the Arfak Mountains in the Vogelkop which is now 
accepted as the type locality of the species (Mayr 1941, 1962, Gilliard 
1969). A single early mounted specimen in the Muséum National 
d’Histoire Naturelle, Paris, labelled as from Amberbaki (north of the 
Arfak Mountains, to the south of the Tamrau Mountains), was 
probably purchased there from local people and was not at its place of 
origin. Ripley (in Mayr & Meyer de Schauensee 1939) and Gilliard (in 
Gilliard & LeCroy 1970) did not record P. carunculata on the Tamrau 
Mountains of the Vogelkop. Rothschild & Hartert (1911) described 
short-tailed P. brevicauda from an adult male collected on Mt. Goliath, 
Oranje Range (eastern Irian Jaya) by A. S. Meek. This simple pattern 
of allopatry was, however, complicated by Ogilvie-Grant’s (1913, 1915) 


C. B. Frith & D. W. Frith 39 Bull BOG. VOL AI (1) 


description of a supposed intermediate species P. intermedia from the 
Utakwa River, below Mt. Carstensz, Snow Mountains, at the western 
end of the central cordillera. ““P. intermedia’’ was established on three 
specimens, consisting of the type, a subadult male (its only adult 
plumage being the central pair of jet-black tail feathers), an immature 
male and a nestling (with nest; see Frith 1970). The characters said to 
distinguish P. intermedia were that it resembled P. carunculata but was 
smaller with a shorter tail, and its wattles were all.lemon-yellow 
(without blue or orange-red). Since then only one other individual bird 
has been attributed to intermedia, this being described as an adult male 
from the Ilaga Valley, Irian Jaya, collected by Ripley (1964). 

While almost all subsequent authors accepted intermedia without 
question as a valid taxon, its position within Paradigalla has proved 
contentious. Most authors agree that intermedia does not constitute 
a valid species but disagree as to which of P. carunculata and 
P. brevicauda it belongs subspecifically. Notable exceptions were 
Rothschild (1921), Mathews (1930), Junge (1939) and Mayr (1941) who 
considered intermedia invalid at any level, and treated it as a synonym of 
P. brevicauda. Gyldenstolpe (1955) found that the material he examined 
was inadequate for decision. Mayr (1941) tentatively considered P. 
brevicauda conspecific with P. carunculata, and this treatment was 
followed by Mayr & Gilliard (1954) and Gyldenstolpe (1955). Iredale 
(1950), Mayr (1962), Frith (1970) and Frith & Frith (1992) treated 
intermedia as a subspecies of P. brevicauda, while Rothschild (1931), 
Ripley (1964), Rand & Gilliard (1967) and Cooper & Forshaw (1977) 
kept it subspecifically under carunculata. While acknowledging P. c. 
intermedia as valid Gilliard (1969) inexplicably detailed the nest and 
nestling collected at the type locality under his account of P. brevicauda. 

With the statement that males and females (although no female 
specimen exists) have a “‘moderately long tail; male with large wattle at 
the base up the upper mandible and forehead and at the base of the 
lower mandible lemon-yellow’’, P. intermedia was resurrected as a full 
species by Cracraft (1992) “‘based on differences in wattle color (e.g. 
Ogilvie-Grant, 1915: 26; Ripley, 1964: 48)’’. 

Two individual paradigallas recently sighted to the south, in the 
Fakfak Mountains, Bombarai Peninsula, Irian Jaya are described as 
having pale yellow facial wattles and swollen pale blue lower mandible 
wattles but without orange-red beneath them, and as having relatively 
short and square-tipped tails (Gibbs 1994). 

No attempt has ever been made to review available series of 
Paradigalla specimens comprehensively with the aim of resolving the 
status and distribution of component taxa. This study seeks to redress 
this situation by using external morphology, biometrical data, and 
zoogeographical patterns. 


Methods 


All skins of Paradigalla spp. were examined in most major museum 
collections world-wide; and CBF measured as many as possible of each 
sex and age class of specimens with a recorded locality. Wing length 


C.B. Fnth © D. W. Frith 40 Bull. BLOC) 1997 Tira 


was taken as the flattened and straightened, thus maximised, chord, 
using a stopped steel rule. The length of the central pair of, and also 
that of the remainder of, tail feathers was measured with a small steel 
rule from the point of entry of the central pair into the bird’s skin to the 
tip of the longest central and remaining feathers. Bill length was 
measured from the bill-tip to the cranio-facial hinge which in 
Paradigalla spp. is markedly high on the forehead toward the level of 
the anterior edge of the eye. Bill width was measured at the anterior 
edge of the nostril. Tarsus length was that of the tarsometatarsal bone. 
All wing and tail measurements were made with the same rules and all 
others with the same electrical digital calipers. 

Of the two taxa that we recognise herein, 50 specimens of P. 
carunculata and 101 of P. brevicauda were examined, and most 
measured, at the following institutions: American Museum of Natural 
History, New York (7/51), Ryksmuseum van Natuurliyke Historie, 
Leiden (4/15), Muséum National d’Histoire Naturelle, Paris (16/0), 
British Museum of Natural History (5/6), Zoologisches Museum, 
Berlin (3/4), Australian Museum, Sydney (0/5), Royal Ontario 
Museum, ‘Toronto (1/4), Staatliches Museum ftir Naturkunde, 
Stuttgart (4/0), Naturmuseum Senckenberg, Frankfurt (3/1), Zoologi- 
cal Museum, Bogor (0/3), Naturhistoriska Riksmuseet, Stockholm 
(0/3), Field Museum of Natural History, Chicago (1/2), Australian 
National Wildlife Collection, CSIRO, Canberra (1/1), Zoologisches 
Museum, Hamburg (0/2), Peabody Museum, Yale University, New 
Haven (2/0), Museum of Comparative Zoology, Berkeley (0/2), 
Museum Koenig, Bonn (1/0), Zoologisk Museum, Kobenhayn (1/0), 
Liverpool Museum and Art Gallery, Liverpool (1/0), Staatliches 
Museum fiir Tierkunde, Dresden (0/1), Papua New Guinea National 
Museum & Art Gallery, Port Moresby (0/1). 

Adult male paradigallas acquire a deep jet-black plumage with areas 
of rich iridescence and velvety sheens and as such are easy to identify in 
the hand, as are subadult males which show some of this feathering 
appearing within the duller plumage of immaturity. Adult females do 
not become as jet-black as adult males, however, and their subadults 
are therefore somewhat more difficult to identify. This must be kept in 
mind when assessing the measurements of specimens presented, there 
being greater variation in the tail length of females, as younger birds 
have shorter (P. carunculata) or longer (P. brevicauda) tails than their 
respective adults. 


Results 


Table 1 presents a summary of mean measurements of paradigalla 
specimens for each sex and age class and for the size differences 
between each sex overall. These results demonstrate that adult male 
and female P. carunculata have a wing length that is on average 15% 
and 9% and a total tail length 60% and 46% longer than those of P. 
brevicauda respectively. Within each sex and age class these differences 
are stressed by mostly mutually exclusive ranges in size, conspicuously 
so in the wing, tail and tarsus lengths of adult males. 


C. B. Frith & D. W. Frith 


41 


TABLE 1 


Bull. B.O.C. 1997 117(1) 


Measurements (mm) of paradigalla skin specimens from various museums 


P. carunculata 
Adult males 


Subadult male 


Immature males 


‘Total males 


Adult females 


P. intermedia 
Males 


P. brevicauda 
Adult males 


Subadult males 


Immature males 


Total males 


Adult females 


Immature females 


Total females 


Wing 


186 
180-201 
6.06 

10 


178 


23 
158 


150 
144-154 
2.96 

26 

153 
152-153 
0.71 

2 


150 


Total tail Central tail 


160 
132-170 
10.60 
10 


137 


133 
121-148 
10.03 


‘Tarsus 


43.1 
38.0-44.5 
2.31 

8 


meee 


43.8 

42 A—44.5 
0.80 

6 


43.4 


42.2 
39:0—44-.0 
1.57 

10 


44.2 
39.6-48.9 
0.93 

25 


A Sal 
40-1—45.3 
2.69 

3 


44.4 
40.6-48.0 
2.15 

20 


45.1 


44.4 
42.1-47.9 
1.49 

25 


44.7 
44.1-45.2 
0.78 

2 


44.4 


Bill length Bull width 


AAW 
co | 
os 
“I 


WhNHn UF 
© | 
oO 
oN 


3 a 
oN 


COUD RF AOUMN NM COM 
(os 


Ni ROS) NY OO Uo ce 
wn | | 
D Ov 
un on 


WwW 


M NOAH poum un ynowun # 
| 
a 
Ww 


No) 


Note: Figures for each age class are (top to bottom) mean, range, SD and sample size. 


C. B. Frith & D. W. Frith 42 Bull. B.O.C. 1997 117(1) 


For adult males and females the average total tail length as a 
percentage of wing length is 71% and 76% respectively in P. carunculata 
and only 34% and 45% respectively in P. brevicauda, while the tarsus 
length as a percentage of wing length is nearly the same (c. 27%) in both 
species. 

On average, adult male P. carunculata have a wing 11% longer than 
their adult females whereas wing length of adult male P. brevicauda is 
only 5% longer than that of adult females of their species. Average 
overall tail length of adult male P. carunculata is 5% longer than of 
females, whereas that of adult male P. brevicauda is, to the contrary, 
22% shorter than that of females. In both sexes and all age classes of 
P. carunculata the central pair of tail feathers are on average 11% longer 
than the remainder of the tail whereas in P. brevicauda the central tail 
feathers are 4% shorter than the rest of the tail. Relative to its much 
larger body size, P. carunculata has a proportionately smaller bill than 
P. brevicauda (‘Table 1). 

One other noteworthy biometric result is that in P. carunculata 
younger individuals have a shorter and less graduated tail than adults 
(contra Ogilvie-Grant 1913, 1915 & Gyldenstolpe 1955) whereas in 
P. brevicauda younger birds have a considerably longer tail than adults. 
Thus the two paradigalla species are distinctly different in size, relative 
proportions and in sexual dimorphism of these characters (contra 
Gilliard 1969, Cooper & Forshaw 1977 for the most part). 

The three immature to adult specimens of supposed intermedia are all 
sexed as male. Of these the BMNH holotype (1916.5.30.1072) is 
subadult and the paratype (1916.5.30.1073) is immature. The third, 
Yale specimen (75320), recorded by Ripley (1964), has black adult 
plumage. Its total tail length of 107mm is (8mm) shorter than 
shortest-tailed adult male P. carunculata and (10mm) longer than 
longest-tailed adult male P. brevicauda. ‘Total tail lengths of the other 
two intermedia specimens are 95 and 94mm, which fall within the 
range of that of young male P. brevicauda and are far (20 mm) shorter 
than those of male P. carunculata of any age. 

‘Table 1 demonstrates that in young male P. brevicauda the central 
rectrices may be longer than the rest of the tail, and in several 
immatures we found them to be up to 5 or 6 mm longer than all other 
tail feathers. A shorter pair of central tail feathers notwithstanding, the 
overall shape of undoubted P. brevicauda individuals may be 
graduated. For example, the lengths of individual tail feathers, from the 
central to the outermost, were 95, 97, 97, 95, 88, 79 in BMNH 
immature/subadult male 1949.62.22 from Tomba, Papua New Guinea. 
This is a more distinctly graduated tail than is that of the type specimen 
of intermedia, which measures 75, 95, 94, 93, 87, 81 respectively. 

In the BMNH holotype the central pair of tail feathers are the only 
jet-black ones of adult plumage and, being unsheathed and evidently 
fully grown (contra Ripley 1964), are shorter (75 mm) than the rest of 
the tail (95 mm) which is only slightly rounded, or graduated. ‘Thus 
this bird would have acquired the shorter tail of P. brevicauda with its 
subsequent moult. Its slightly graduated tail, which would probably 
have become even more truncate with the subsequent moult, is 


C. B. Frith & D. W. Frith 43 Bull. B.O.C. 1997 117(1) 


identical to that of several undoubted subadult male P. brevicauda (e.g. 
BMNH 1969.41.806) from Mt. Kunupi in the central cordillera of 
Irian Jaya. 

No specimens of intermedia show any sign of the additional bare 
orange-red area of skin (of P. carunculata) beneath the blue lower 
mandible wattle. Noteworthy too is that the colours of facial bare parts 
were not recorded for the holotype of intermedia. These are noted on 
the label of the paratype, however, as ‘“‘nasal flaps base of bill 
lemon-yellow’’. In both of these specimens, however, the dried upper 
facial wattles are now a mustard-yellow while the lower ones are 
contrastingly blackish. Of the third intermedia specimen, at Yale, 
Ripley (1964) recorded all facial wattles of the freshly dead bird as clear 
lemon yellow. While describing this individual as an adult male, Ripley 
recorded on its label that its gonads were unenlarged at a time (10 
September) when an adult might be expected to be in breeding 
condition. The tail of the latter specimen is graduated but that of the 
holotype and paratype is barely so, being merely slightly ‘rounded’ (see 
Discussion). 

A point requiring clarification here relates to the unsexed nestling 
(BMNH 1916.5.30.1074) taken with its nest at the Utakwa River in 
conjunction with the collection of type material of intermedia. Vague 
wording by some authors subsequent to Ogilvie-Grant (1913, 1915) 
may give the impression that the male holotype or paratype of 
intermedia was collected with the nestling and nest. In fact the nestling 
was collected at Camp 9 on 27 January and the two male birds at Camp 
6c on 24—25 February. Moreover, in view of what we now know about 
the nesting biology of P. brevicauda, and the fact that P. carunculata 
appears to have been a putative parent of hybrids involving other 
bird of paradise species, it would seem most unlikely that male 
P. “intermedia” would attend nests (Frith & Frith 1992). 

The distributions of paradigalla taxa, derived from an examination 
of specimens and the literature, shown in Figure 1 demonstrate that 
P. carunculata is confined to the Arfak Mountains of the eastern 
Vogelkop proper. The species P. brevicauda is recorded from the 
central mountain ranges of New Guinea from the Mt. Wilhelm and 
Mt. Karimui area of Papua New Guinea westward to the Wissel lakes 
area, Weyland Mountains of Irian Jaya. 


Discussion 


It might be thought that the three male intermedia skins reflect a 
tendency for males of P. brevicauda to have longer tails at the western 
limits of the species range (Fig. 1). This is not the case, however, as six 
birds further to the west, on the Weyland Mountains, have the typically 
- short tail of P. brevicauda. These specimens, AMNH 0049951, 302983, 
678340, 678348, 302981 and 678347, have tail lengths of 44, 45, 45, 49, 
51 and 52mm respectively, all shorter than the average adult male 
P. brevicauda tail length (Table 1). Accordingly, we consider the three 
specimens of free-flying “intermedia”? to be relatively young, and 
therefore long-tailed, individuals of P. brevicauda of which only one has 


C. B. Frith @ D. W. Frith 44 Bull. B.O.C. 1997 117(1) 


Figure 1. Map of mainland New Guinea showing the border between Irian Jaya (IJ) and 
Papua New Guinea (PNG) and the distribution of the two Paradigalla species based 
on collected specimens and published field sightings that are considered reliable. Note: 
Solid circles=P. brevicauda; circles with stars within=P. (intermedia) brevicauda; open 
circles=P. carunculata; ?=P. sp. sighted in Fakfak Mountains (see text). Note: a single 
circle may encompass several collecting locations. Locations of some key features are 
indicated by (left to right): V=Vogelkop; large F=Fakfak; W=Weyland Mts.; 
U=Utakwa River; G=Mt. Goliath; F=Fly River; H=Mt. Hagen; M=Madang; L=Lae; 
P=Port Moresby. 


a tail length (see below) greater than recorded in that species (Table 1). 
In particular the central tail feather length of the holotype of intermedia 
(75 mm) is far more compatible with that of P. brevicauda than with 
P. carunculata (Table 1). Moreover, we are able to confirm Mayr & 
Gilliard’s (1954) comment that specimens of P. brevicauda from the 
extreme eastern part of the species range (Mt. Hagen and Bismarck 
Range) are indistinguishable from those from the extreme west 
(Weyland Mountains). 

Given that we found that the dried upper facial wattles (lemon yellow 
in life) were a mustard-yellow but the lower ones (blue in life) were 
blackish in all skins of adult P. brevicauda, the differences in wattle 
colours of live birds are clearly reflected in dried skins. That the upper 
wattle was mustard-yellow and the lower one blackish in the zntermedia 
type material strongly suggests that, if not blue in life, the lower wattle 
was to have become so. T'wo immature BMNH P. brevicauda 
specimens (1949.62.22 and 1969.41.806 from Tomba, Papua New 
Guinea and Mt. Kunupi, Irian Jaya, respectively) have lower wattles 
mustard-yellow as their upper ones but with darker blackish pigment 
apparent along their upper and lower edges and at their posterior end. 
These immature birds thus appear to suggest that the blue wattle 
colour only begins to be acquired by advanced immatures or subadults. 
This is supported by the fact that the dried wattles in the BMNH 
nestling specimen are pale mustard-yellow throughout, with no sign of 
dark pigment. A near-fledged large nestling photographed in the wild 
also had wattles dull lemon yellow throughout (Frith in Coates 1990). 


C. B. Frith & D. W. Frith 45 Bulls OG. 1997, 1de7 (4) 


Moreover, Gilliard & LeCroy (1961) described a freshly dead subadult 
(‘virtually adult’) male collected at Mt. Ifal, Victor Emanuel 
Mountains as having the upper facial wattle bright lime yellow and the 
lower ‘gape’ wattle “‘dull yellowish grey, with black contour lines 
below’. The latter feature matches the condition noted in a nestling 
and in dried specimens of P. brevicauda we consider to be subadults as 
detailed above. Thus, the supposed distinctive character of an 
all-yellow facial and bill wattle cannot be accepted as taxonomically 
diagnostic of intermedia, as it occurs in young and subadult P. 
brevicauda. We consider it quite possible that young P. carunculata will 
also lack blue colour in the lower mandible wattle. The additional area 
of distinctly orange-red bare skin beneath the base of the blue wattle of 
P. carunculata appears diagnostic of the species, although this area has 
been described as bare but yellow in a single freshly dead adult male 
P. brevicauda on Mt. Ifal, Victor Emanuel Mountains (Gilliard & 
LeCroy, 1961). 

In attributing his Ilaga Valley specimen to P. c. intermedia, Ripley 
(1964) supported his case by presuming that the shorter central tail 
feather pair of the holotype of intermedia must have been ‘“‘not fully 
moulted out’; but these feathers are in fact fully grown in that 
specimen. Ripley also made much of the fact that the tail of his own 
specimen “has the normal wedge shape of carunculata, the central 
rectrices being the longest’. Our measuring of this specimen, however, 
gave a total tail length of 107 mm, with the central pair of rectrices one 
millimetre shorter at 106mm. All three intermedia specimens in fact 
have the central pair of rectrices shortest whereas in all male (adult and 
immature) P. carunculata specimens (n=16) the central pair are 
conspicuously longest. The tail lengths of the two BMNH specimens 
of intermedia (94 and 95 mm) fall within the range of younger male 
P. brevicauda (‘Table 1) but that of Ripley’s specimen (107 mm) is 
longer than any male P. brevicauda but shorter than any P. carunculata. 
Notwithstanding the black adult-like plumage of Ripley’s specimen, we 
interpret its exclusively yellow wattles and its just shortest central 
rectrices as indicative of a relatively young individual P. brevicauda in 
first adult plumage retaining the longer (longest known) tail typical 
of immaturity. That Ripley’s bird had unenlarged gonads on 10 
September permits the possibility it is in fact a less than fully sexually 
mature P. brevicauda which could account for its long tail (see above). 
That it had a graduated tail, moreover, does not exclude it from 
P. brevicauda as individuals of the latter are known with graduated tails 
(see Results). Significantly, the wing length of Ripley’s specimen 
(155mm) is not intermediate between that of P. carunculata and 
P. brevicauda but is shorter than all P. carunculata specimens (n=28) 
and is in fact short for male P. brevicauda (‘Table 1). 

The above facts show that, in general size and shape, the specimens 
of supposed intermedia are in fact far closer to P. brevicauda than to 
P. carunculata, as emphasised by Mathews (1930) and Mayr (1941, 
1962). In addition, the supposed diagnostically yellow lower wattle of 
intermedia is shown here to be characteristic of immature/subadult 
P. brevicauda and probably also P. carunculata; and this correlates with 


C. B. Fnth & D. W. Frith 46 Bull. B.O.C. 1997 117(1) 


a longer tail in younger birds. Contrary to P. brevicauda, immatures of 
P. carunculata have a shorter tail than adults (contra Gyldenstolpe 
1955: 135). For these reasons, and the fact that “‘intermedia’’ is known 
only from within the geographic range of brevicauda, not outside and 
adjacent to carunculata (Fig. 1), the taxon intermedia should be 
considered invalid. 

In overall and proportionate wing and tail lengths, which differ 
sexually from P. brevicauda, the Arfak paradigalla is most distinct. On 
these characters alone P. carunculata and P. brevicauda may be judged 
good species, particularly as we found no clinal trend towards longer 
tails in western populations of P. brevicauda. Until such time as 
convincing evidence suggests otherwise, the four birds presently known 
as intermedia should be considered as representing immature and/or 
subadult individuals of monotypic P. brevicauda. 'The recent resur- 
rection of P. intermedia as a full species (Cracraft 1992), confined to the 
Carstensz Peak type locality area entirely within the extensive range of 
typical P. brevicauda populations, is zoogeographically questionable, 
quite aside from the strongly contradictory morphological and 
biometrical evidence presented here. 

P. carunculata, as far as is known, is endemic to the Arfak 
Mountains, with a congeneric representative elsewhere on the core 
cordilleras of New Guinea, as in several other west Irian Jayan montane 
species such as the Vogelkop Scrub-wren Sericornis rufescens, 
Grey-banded Mannikin Lonchura vana, Vogelkop Bowerbird Ambly- 
ornis inornatus, Arfak Astrapia Astrapia mgra and Western Parotia 
Parotia sefilata (Beehler et al. 1986). 

The identity of two paradigallas sighted in the Fakfak Mountains, 
Bomberai Peninsula, with pale yellow upper and blue lower facial 
wattles, without any orange-red skin, and square-tipped short tails 
remains problematical (Gibbs 1994). On zoogeographical grounds, the 
Fakfak birds would appear most likely to be of P. carunculata stock 
(Beehler et al. 1986). While a markedly short and square-ended tail as 
described would suggest P. brevicauda (see ‘Table 1), the drawing 
published by Gibbs could be of relatively short-tailed individuals of 
P. carunculata. If the birds seen by Gibbs were in fact P. carunculata, 
they would most probably have been young females, given their 
relatively short tails (Table 1). Because Gibbs (1994) also presents 
evidence suggesting that at least two or three distinctly new and 
unknown forms of passerines apparently await formal discovery and 
description from that region, the collection of paradigallas there is 
needed urgently. 


Summary 


The New Guinea montane genus Paradigalla (Paradisaeidae) is reviewed, based on an 
examination of most specimens in major world museums, and the literature. One 
hundred and fifty specimens were measured; their biometrics are presented and 
discussed, and their locations are mapped and presented. Only two taxa are accepted: 
larger and longer-tailed P. carunculata confined to the Arfak Mountains of the Vogelkop, 
Irian Jaya, in which adults have yellow, blue and red facial wattles and young birds a tail 
shorter than adults; and smaller and shorter-tailed P. brevicauda throughout mountains 
of the main trunk of New Guinea, in which adults lack the red facial wattle and young 


C. B. Frith & D. W. Frith 47 Era. #5 OC. T9971 19 (1) 


birds have a tail longer than adults. The four known specimens of P. c. intermedia were 
found to show no distinctive characters and on morphological, biometrical and 
zoogeographical evidence are interpreted as relatively young individuals of monotypic 
P. brevicauda. 


Acknowledgements 


For much appreciated permission to study the ornithological collections in their care we 
sincerely thank the following persons and institutions: Dean Amadon, Chris Blake, 
Walter Bock, Joel Cracraft, Mary LeCroy, Manny Levine and Lester Short, American 
Museum of Natural History, New York; Robert Prys-Jones, Peter Colston and Michael 
Walters, British Museum of Natural History; Ned Johnson, Museum of Vertebrate 
Zoology, University of California, Berkeley; David Willard, The Field Museum, 
Chicago; Raymond A. Paynter Jr., Museum of Comparative Zoology, Cambridge; James 
Dick and Brad Millen, Royal Ontario Museum, Toronto; Fred C. Sibley, Peabody 
Museum, Yale University; Richard Schodde, Ian Mason and John Wombey, Australian 
National Wildlife Collection, Canberra; Walter Boles and Wayne Longmore, Australian 
Museum, Sydney; Mohammad Amir and Darjono, Museum Zoologicum Bogoriense, 
Bogor; Frank Bonaccorso, Ilaiah Bigilale and Paul Wanga, National Museum and Art 
Gallery of Papua New Guinea, Port Moresby; Clem Fisher, National Museums & 
Galleries, Liverpool; Rene Dekker and Peter Van Dam, Rijksmuseum van Natuurlijke 
Historie, Leiden; Per Ericson, Gunnar Johansson, Gé6ran Frisk, SMNH, Stockholm; Jon 
Fjeldsa, Zoologisk Museum Kgobenhavns Universitet, Kobenhaven; Eric Pasquet, 
Muséum National d’Histoire Naturelle, Paris; Siegfried Eck, Staatliches Museum ftir 
Tierkunde, Dresden; Prof. B. Stephan, Curator of Birds, Zoologisch Museum, Berlin; 
R. van den Elzen, Museum Alexander Koenig, Bonn for kindly forwarding skins to 
Frankfurt; Claus Konig, Curator of Birds, Staatliches Museum ftir Naturkunde, 
Stuttgart for kind hospitality in addition to access to collections; D. S. Peters, Prof. Dr 
Steinbacher, Martina Kiisters and Karin Béhm, Forschungsinstitut und Naturmuseum 
Senckenberg, Frankfurt; H. Hoerschelmann, Zoologisches Institut und Zoologishes 
Museum, Hamburg for kindly sending skins to Frankfurt. 

We particularly thank Mary LeCroy for remarkably kind hospitality in New York, and 
the Chapman Fund of the American Museum of Natural History for support. Bruce 
Beehler, Brian Coates, Jared Diamond, Mary LeCroy and Richard Schodde improved 
this contribution by constructively commenting on an early draft. 

Dedicated to Prof. Dr Ernst Mayr whose monumental work on birds of paradise, other 
New Guinea birds and very much more is inspirational, and who long ago correctly 
placed P. intermedia into the synonymy of P. brevicauda. 


References: 

Beehler, B. M., Pratt, T. K. & Zimmerman, D. A. 1986. Birds of New Guinea. Princeton 
Univ. Press. 

Coates, B. J. 1990. The Birds of Papua New Guinea. Vol. 2. Dove, Alderley, Queensland. 

Cooper, W. T. & Forshaw, J. M. 1977. The Birds of Paradise and Bower Birds. Collins, 
Sydney. 

Cracraft, J. 1992. The species of the birds-of-paradise (Paradisaeidae): applying the 
phylogenetic species concept to a complex pattern of diversification. Cladistics 8: 
1-43. 

Frith, C. B. 1970. The nest and nestling of the Short-tailed Paradigalla Paradigalla 
brevicauda (Paradisaeidae). Bull. Brit. Orn. Cl. 90: 122-124. 

Frith, C. B. & Frith, D. W. 1992. Nesting biology of the Short-tailed Paradigalla 
Paradigalla brevicauda. Ibis 134: 77-82. 

Gibbs, D. 1994. Undescribed taxa and new records from the Fakfak Mountains, Irian 
Jaya. Bull. Brit. Orn. Cl. 114: 4-12. 

Gilliard, E. T. 1969. Birds of Paradise and Bower Birds. Weidenfeld & Nicolson, 
London. 

Gilliard, E. T. & LeCroy, M. 1961. Birds of the Victor Emanuel and Hindenburg 
Mountains, New Guinea. Results of the American Museum of Natural History 
Expedition to New Guinea in 1954. Bull. Am. Mus. Nat. Hist. 123: 1-86. 


R. }. Dowsett G A. Moore 48 Bulln~B.O.C, 1997 ALG) 


Gilliard, E. T. & LeCroy, M. 1970. Notes on birds from the Tamrau Mountains, 
New Guinea. Am. Mus. Novit. 2420. 

Gyldenstolpe, N. 1955. Notes on a collection of birds made in the Western Highlands, 
Central New Guinea, 1951. Ark. Zool. Series 2, 8: 1-181. 

Iredale, T. 1950. Birds of Paradise and Bowerbirds. Georgian House, Melbourne. 

Junge, G. C. A. 1939. The Birds of South New Guinea. Part 2, passerines. Nova Guinea 
3: 1-94. 

Lesson, R.-P. 1835. Histoire naturelle des oiseaux de Paradis et des épimaques. Paris. 

Mathews, G. M. 1930. Systema Avium Australasianarum, a systematic list of the birds of 
the Australasian region. Part 2. British Ornithologists’ Union, London. 

Mayr, E. 1941. List of New Guinea Birds. American Museum of Natural History, 
New York. 

Mayr, E. 1962. Family Paradisaeidae. In: E. Mayr & J. C. Greenway, Jr. (eds), Check-list 
of Birds of the World. Vol. 15. Museum of Comparative Zoology, Harvard. 

Mayr, E. & Meyer de Schauensee, R. 1939. Zoological results of the Denison-Crockett 
South Pacific Expedition for the Academy of Natural Sciences of Philadelphia, 
1937-1938. Part IV.—birds from northwest New Guinea. Proc. Acad. Nat. Sci. 
Philadelphia 91: 97-144. 

Mayr, E. & Gilliard, E. T. 1954. Birds of central New Guinea. Results of the American 
Museum of Natural Expedition to New Guinea in 1950 and 1952. Bull. Am. Mus. 
Nat. Hist. 102: 315-374. 

Ogilvie-Grant, W. R. 1913. [Paradigalla intermedia, sp.n.]|. Bull. Brit. Orn. Cl. 31: 
99-106. 

Ogilvie-Grant, W. R. 1915. Report on the birds collected by the British Ornithologists’ 
Union Expedition and the Wollaston Expedition in Dutch New Guinea. Ibis, Jubilee 
Supplement 2. 

Rand, A. L. & Gilliard, E. T. 1967. Handbook of New Guinea Birds. Weidenfeld & 
Nicolson, London. 

Ripley, S. D. 1964. A systematic and ecological study of birds of New Guinea. Bull. 
Peabody Mus. Nat. Hist. 19: 1-87. 

Rothschild, Lord. 1921. On some birds from the Weyland Mountains, Dutch New 
Guinea. Novit. Zool. 28: 280-294. 

Rothschild, Lord. 1931. On a collection of birds made by Mr. F. Shaw Mayer in the 
Weyland Mountains, Dutch New Guinea, in 1930. Novit. Zool. 36: 250-276. 

Rothschild, W. & Hartert, E. 1911. Preliminary descriptions of some new birds from 
central New Guinea. Novit. Zool. 18: 159-161. 


Address: C. B. and D. W. Frith, P.O. Box 581, Malanda, Nth Qld 4885, Australia. 


© British Ornithologists’ Club 1997 


Swamp warblers Acrocephalus gracilirostris 
and A. rufescens at Lake Chad, Nigeria 


by R. F. Dowsett GS Amberley Moore 
Received 30 December 1995 


This paper aims to clarify the status of two swamp warblers at Lake 
Chad, the Lesser Swamp Warbler Acrocephalus gracilirostris and the 
Greater Swamp Warbler A. rufescens. 

While living at Malamfatori (13°37’'N, 13°20’E) on the Nigerian 
shore of Lake Chad during 1968, R.J.D. noted A. gracilirostris in 
song from the time of his arrival at the end of February. This species 


R. ¥. Dowsett & A. Moore 49 Bull. B.O.C. 1997 117(1) 


was fairly often in small numbers in lakeshore Cyperaceae and 
Phragmites in the vicinity of Malamfatori, but A. rufescens only 
rarely. The latter was normally confined to extensive areas of Papyrus 
elsewhere on the lake, such an ecological distinction being normal 
in parts of Africa where the two are sympatric. At Malamfatori in 
May 1968 R.J.D. ringed and netted four A. gracilirostris (and two 
A. rufescens) and has retained detailed notes of two of the A. 
gracilirostris. The five Lesser Swamp Warblers mentioned (under the 
name Calamoeceter leptorhyncha) by Sharland (1969) in the ringing 
report for Nigeria, ringed to the end of 1968, presumably included 
these four birds netted at Malamfatori. Each of the two ringed by 
R.J.D. had a wing-length of 69 mm, and their weights were 12.3 and 
12.7 g. Other measurements included tarsus 27-28 mm and tail 
63-65 mm. These data accord well with those of six specimens of 
A. gracilirostris collected at Lake Chad by Boyd Alexander, and 
re-examined by Bates (1930). They also fall well within the range of 
A. gracilirostris at Jekara (Aidley & Wilkinson 1987) and that of 131 
A. g. leptorhynchus ringed by R.J.D. (unpubl.) in southern Zambia: 
wings 58—74 (mean 67 mm) and weights 11.3—20.3 (mean 14.8 g). The 
weight data for the Greater Swamp Warbler at Jekara showed it to be 
50% heavier than A. gracilirostris, with no overlap. In another area of 
sympatry in Kenya Britton (1978) also gave mean wing-lengths and 
weights of A. rufescens that were considerably greater than those of 
A. gracilirostris. 

The Lesser Swamp Warbler is also listed by Dowsett (1993: 106) 
from Chad, on the authority of Salvan (1968). But it must be admitted 
that Salvan himself had as his only certain records of the two swamp 
warblers the type specimens collected by Alexander. 

Through the kind cooperation of the Federal Fisheries research 
staff R.J.D. was able to make three voyages, each of three days, on 
Lake Chad in the boat “El Kanemi’’, between late June and early 
August 1968. He found both warblers in Chadian territory, with 
A. gracilirostris especially numerous locally. Separation of the two in 
the field was based mainly on the notably larger size and longer tail of 
A. rufescens; although the latter does tend to have a deeper voice, this 
distinction is not always evident (Dowsett-Lemaire 1994). Vielliard 
(1972) also found A. gracilirostris to be abundant on the Chad side 
of the lake, collecting no fewer than 22 specimens (compared to 4 
A. rufescens, which he did not consider to be rare). We have no definite 
record from the Niger sector of the lake, although R.J.D. did notice 
that around Nguigmi (Nguimi) there was suitable habitat in 1968. The 
species presumably does not occur there now, with the retreat of the 
surface area of Lake Chad. 

Alexander (1908) gave the locality for the type specimen of A. 
gracilirostris neglectus merely as Lake Chad, and the date as 5 December 
1904. However, A.M. has been able to determine a more exact 
type-locality, from consideration of the details in Alexander’s field 
notebooks (held in the Natural History Museum at Tring), together 
with the account of the Alexander-Gosling expedition to the Lake Chad 
area published by Alexander (1907a). Boyd Alexander and P. A. Talbot 


R. F. Dowsett & A. Moore 50 Bull. B.O.C. 1997 117(1) 


began to survey and map part of Lake Chad in December 1904. A map 
published by Alexander (op. cit.) is marked with their route and a 
number of dates. Correlation of the dates and places with those given in 
Alexander’s notebook provides an itinerary which is likely to be 
reliable. 

Alexander’s notebook contains for each bird collected his collector’s 
number, the date and place of collection and some field observations. 
Warren & Harrison (1971) did not give the collector’s numbers for 
these holotypes, nor are they in the BM register for the Alexander 
Collection (Mrs F. E. Warr pers. comm. to A.M.). The type specimen 
of A. g. neglectus in the BM bears Alexander’s label with his collector’s 
number 406; the entry in his notebook for no. 406 reads ‘“‘Calamocichla 
¢ Lake Chad 5.12.904 First observed in the Maria bushes. Breeding. 
Song like that of C. brevipennis ...’”’. 

From 30 November 1904 Alexander worked the Yo area, from 
Kopichi (near Kukawa 12°55’N, 13°35’E), visiting both sides of the 
Yobe basin. He commented that the villages on the islands were cattle 
stations, inhabited for short periods only. The expedition did not cross 
to the eastern (Chad) shore of the lake until 10/11 December 1904. 
‘These dates accord well with those published for the various type 
specimens collected by Alexander during 1904—05 (Dowsett in prep.), 
November localities such as Kukawa and Yo being in the Nigerian 
sector of the lake and within a 40 km radius of Malamfatori. We would 
therefore clarify the type-locality of A. gracilirostris neglectus as ‘““Lake 
Chad, Nigeria’’. 

Alexander (1907b) did not give data for his type specimen of 
A. rufescens chadensis; its label bears the date 19 May 1905 and locality. 
This is given as ‘““Wurmda”’ in Warren & Hastings (1971), but in 
Alexander’s notebook and elsewhere written ““Wunnda’’. The type 
specimen is also held in the BM and bears his label with his number 
1094. His notebook records “‘1094 Calamocichla ¢ Wunnda 19.5.1905 
Found in the depth of thick maria clumps. First time observed on 
Karraragga’’. From Alexander’s published map Wunnda (13°30'N, 
14°25’E) and Karraragga (13°22’N, 14°21’E), an area of “‘acacia, good 
pasture’’, are situated on the north shore of the eastern bay of the lake, 
in Chad. 

Elgood et al. (1994) reported only one definite record of A. 
gracilirostris in Nigeria, from Jekara in Kano District (Wilkinson & 
Aidley 1982). At Jekara dam Aidley & Wilkinson (1987) caught and 
ringed 46. The Lesser Swamp Warbler was omitted from the first 
Nigerian checklist (Elgood 1981) through an oversight; it had been 
listed for the Malamfatori area by Hopson (1964), Dowsett & Hopson 
(1969) later adding A. rufescens to the local list. 

Although Elgood et al. (1994) suggest that Greater Swamp Warblers 
in the Nigerian sector of Lake Chad “‘are possibly A. r. chadensis, though 
this has not been proven’’, there is no doubt of the type-locality. The 
question is perhaps rather whether or not chadensis is separable from the 
nominate race, which ranges from Cabinda to southern Nigeria. As 
pointed out by White (1960) the distinctions are slight, and presumably 
the range of the species is for practical purposes continuous. 


R. 3. Dowsett & A. Moore 51 Bull. B.O.C. 1997 117(1) 


Acknowledgements 


We are most grateful to Mrs F. E. Warr and to Michael Walters of the Natural History 
Museum, Tring (BM). Mrs Warr kindly checked the entries in Alexander’s notebooks 
and Mr Walters confirmed the collector’s numbers of the two type specimens. Dr D. J. 
Aidley read the first draft of this note. 


References: 

Aidley, D. J. & Wilkinson, R. 1987. The annual cycle of six Acrocephalus warblers in a 
Nigerian reed-bed. Bird Study 34: 226-234. 

Alexander, B. 1907a. From the Niger to the Nile. Edward Arnold, London. 

Alexander, B. 1907b. [A new species of Calamocichla.| Bull. Brit. Orn. Cl. 19: 63. 

Alexander, B. 1908. [Descriptions of two new species of birds from Lake Chad.] Bull. 
Bat Ors Cl. -23-: 33. 

Bates, G. L. 1930. Handbook of the Birds of West Africa. John Bale, Sons & Danielsson, 
London. 

Britton, P. L. 1978. Seasonality, density and diversity of birds of a papyrus swamp in 
western Kenya. [bis 120: 450-466. 

Dowsett, R. J. 1993. Afrotropical avifaunas: annotated country lists. Tauraco Res. Rep. 5: 
1-322. 

Dowsett, R. J. in prep. Checklist of birds of the Afrotropical and Malagasy regions. Vol. 2: 
Subspecies and type-localities. Tauraco Press, Liége. 

Dowsett, R. J. & Hopson, A. J. 1969. Additions and amendments to the List of birds of 
Malam’fatori, Lake Chad. Bull. Nigerian Orn. Soc. 6: 53-55. 

Dowsett-Lemaire, F. 1994. The song of the Seychelles Warbler Acrocephalus sechellensis 
and its African relatives. Ibis 136: 489-491. 

Elgood, J. H. 1981. The Birds of Nigeria. B.O.U., London. 

Elgood, J. H., Heigham, J. B., Moore, A. M., Nason, A. M., Sharland, R. E. & Skinner, 
N. J. 1994. The Birds of Nigeria. B.O.U., Tring. 

Hall, B. P. & Moreau, R. E. 1970. An Atlas of Speciation in African Passerine Birds. Brit. 
Mus. (Nat. Hist.), London. 

Hopson, A. J. 1964. Preliminary notes on the birds of Malamfatori, Lake Chad. Bull. 
Nigerian Orn. Soc. 1(4): 7-15. 

Salvan, J. 1968. Contribution a létude des oiseaux du Tchad. (Part 4). Ozseau 38: 
249-273. 

Sharland, R. E. 1969. Ethiopian birds ringed in Nigeria and Ghana. Bull. Nigerian Orn. 
Soc. 6: 50-52. 

Vielliard, J. 1972. Données biogéographiques sur |’avifaune de |’ Afrique centrale (suite). 
Alauda 40: 63-92. 

Warren, R. L. M. & Harrison, C. J. O. 1971. Type-specimens of Birds in the British 
Museum (Natural History). Vol. 2 Passerines. Brit. Mus. (Nat. Hist.), London. 
White, C. M. N. 1960. A check list of the Ethiopian Muscicapidae (Sylviinae). Part I. 

Occ. Pap. Nat. Mus. S. Rhod. 24B: 399-430. 
Wilkinson, R. & Aidley, D. J. 1982. Additions to local avifaunas: Kano State. Malimbus 
4: 107. 


Address: R. J. Dowsett, 194 rue de Bois de Breux, B-4020 Jupille, Liége, Belgium. Mrs 
A. M. Moore, 1 Uppingham Road, Oakham, Rutland LE15 6JB, U.K. 


© British Ornithologists’ Club 1997 


P. A. Clancey 52 Bull. B.O.C. 1997 117(1) 


Variation in the Cape Penduline Tit 
Anthoscopus minutus of the southern 
Afrotropics 


by FA Clancey 
Received 4 Fanuary 1996 


The austral African penduline tits Anthoscopus minutus (Shaw), 1812, 
and Anthoscopus caroli (Sharpe), 1871, form a species pair with a 
minimal measure of overlap, with the former disposed in the dry west 
and interior, and carvoli replacing it in the north and east of their joint 
range. While generally viewed as closely allied, their proportions 
suggest they are not of immediate ancestral origin, the tail-length of 
minutus being appreciably longer than in carols (33-37, versus 
27—28.5 mm), which also has the forehead, distal face and chin boldly 
patterned with black, which surfaces are more or less bland in caroli. 
The range of the eastern caroli encompasses a wider range of woodland 
facies than that of the western minutus, whose range comprises a more 
mesic body of scrubland habitat types, being ecologically karroid. 
A. minutus, unlike carol, is restricted to the South West Arid Zone of 
the Afrotropics, whereas its analogue in southern Africa is distributed 
in some six or seven named races as far north as northern Uganda and 
adjacent northeastern Africa, judging from Snow’s (1967) treatment of 
the species in Peters’ Check-list, the most recent assessment. 

A high proportion of specimens, which I take to be in first-year dress, 
are often difficult to place in subspecies, except in the case of the southern 
race gigi, in which a brownish cast to the sides of the body and flanks is 
diagnostic. Birds are in newly assumed dress from about late April to 
early June. Seventy specimens were examined during this research. 

Specimens agreeing with topotypes of A. m. minutus are present along 
the entire west coast of the Cape Province and Namibia from about the 
‘Tropic of Capricorn to the Cape of Good Hope, and are characterized by 
having the throat and breast coloration light neutral grey, this merging 
insensibly into pale yellow (close to Maize Yellow; Ridgway 1912), the 
sides of the body and flanks tending to be on the whole slightly darker. 
Over the upper-parts, the pileum and hind neck differ in being normally 
a little darker than the rest of the dorsal surface. The population present 
in Namibia lying to the north of those just dealt with are on the whole 
lighter in freshly plumaged adults, are starkly whiter over the chin and 
fore-throat, and are strikingly deeper yellow (close to Spectrum Yellow; 
Ridgway 1912) over the rest of the ventral parts. It was to such birds that 
Reichenow (1905) gave the name A. m. damarensis, the type-locality 
designated later as Ovaquenyama, Owambo, by Macdonald (1952). The 
range of damarensis extends eastwards to the northern and western 
aspects of Zimbabwe in the Matopos and to the Transvaal. In the east of 
the range, birds of damarensis-type are found as far south as Kuruman, 
but with the species clearly uncommon in this sector a picture of the 
racial pattern is of necessity tentative. 


P. A. Clancey 53 Bull. B.O.C. 1997 117(1) 


TABLE 1 
Free State and eastern Cape localities from which specimens were found to agree closely 
with paratypical A. m. gigi 


Free State 

Bloemfontein 11 May 1957 
Glen Lyon Farm, Glen 28 April 1960 
Sandymountpark, Fauresmith 22 March 1994 
Cape 

Colesberg 19 August 1952 
Melton Wold, Victoria West 26 May 1989 
Graaff-Reinet 12 July 1967 
‘Tarkastad 1 July 1959, 3 April 1969, 8 September 1969 
Fort Beaufort 28 July 1961 
Oudtshoorn 10 April 1956 
Thomas River, Cathcart 21 May 1964 


Examination of the distribution as defined in the Atlas of Speciation 
by Moreau & Hall (1970) reveals a singular absence of records of the 
Cape Penduline Tit over an extensive swathe of country lying inland 
beyond the coastal strip between the Tropic in the north and the 
territory to the west of Cape Agulhas area in the far south. It is hard 
to concede that this has come about fortuitously or as a result of 
the dearth of specimen records, certainly to the south of the Orange 
R., which has been well covered in recent times by collectors. To 
the immediate east of this major range hiatus lies a small detached seg- 
ment of nominate minutus as well as elements of an allied form which 
Winterbottom (1959) separated, on a small sample of three skins from 
Oudtshoorn, as A. m. gigi, a race which seemingly has been 
consistently ‘overlooked: Snow (1967) treated it as a straight synonym 
of nominate minutus without discussion. The taxon was briefly dealt 
with by me in my treatment of the present penduline tit in the 
S.A.O.S. Checklist, when I was able to study a limited sample. This 
has now been augmented to a series of eighteen skins, as listed in 
Table 1. 

A. m. gigi is found to be a relatively well-marked subspecies 
distinguishable from the nominate form in its darker upper-parts which 
are dark Greyish Olive (Ridgway 1912), the olive-buff of the dorsal 
parts confined to the rump and upper tail-coverts. On the underside, 
the chin and fore-throat are greyish-brown, merging into olivaceous 
buff over the rest of the ventral surface, the sides and flanks overall 
darker. There is no vestige of white or clear yellow to the under-parts 
in what I take to be the definitive plumage of the present race. In the 
case of the two sub-adults from Glen Lyon Farm, these specimens 
differ in having the sides of the venter light brownish, as opposed to 
white or clear yellowish in the other two taxa. 

‘The pattern of variation thus exhibited warrants recognition of three 
well-characterized races, with ranges as indicated in Figure 1. Details 
of each are given below. 


P. A. Clancey 54 Bull. B.O.C. 1997 117(1) 


Figure 1. Sketch-map of southern Africa showing the range of the Cape penduline Tit 
and of its three subspecies: 1, Anthoscopus minutus damarensis Reichenow; 2, Anthoscopus 
minutus minutus (Shaw); 3, Anthoscopus minutus gigi Winterbottom. 


Anthoscopus minutus damarensis Reichenow 

Anthoscopus minutus damarensis Reichenow, Die Végel Afrikas, 
vol. ii, 1905, p.526: Damaraland, restricted to Ovaquenyama, 
Owambo, Namibia, by Macdonald (1952), Bull. Brit. Orn. Cl. 72, 
p. 49. 

Adult with the dorsal surfaces deep olive-buff, the crown and hind 
neck rather more ochreous than the rest of the dorsum; on the 
underside with the chin and fore-throat clear white, merging into 
spectrum yellow medio-ventrally. 

First-year dress with the dorsal surface as described for the adult but 
with the chin and fore-throat less starkly white, merging into more 
dilute yellow below. 

Wing: 55 47-51.5 (49.3), n=9; 99 48-52.5 (50.2) mm, n=11. 
Range. Northern Damaraland from about the Windhoek district to 
northern Owamboland, east to the Caprivi Strip and northeastern 
Botswana, western Zimbabwe in the Matopos Hills district and near 
Bulawayo, and western Transvaal; further south reaching as far as the 
Kuruman district in the northern Cape. Also recorded by Moreau & 
Hall (1970) as occurring on the Angolan desert coast as far north as 


Luanda. Generally confined to areas with annual rainfall less than 
250 mm. 


P. A. Clancey 55 Bull. B.O.C. 1997 117(1) 


Anthoscopus minutus minutus (Shaw) 
Sylvia minuta Shaw, in Shaw & Nodder, Nat. Misc., No. 23, 1812, 
pl. 997: Elephant R., restricted to Heerenlogement, S.W. of Klaver, 
western Cape, by Macdonald (1952), Bull. Brit. Orn. Cl. 72, p. 48. 
Adult differs from damarensts in having the crown and hind-neck dull 
olive-buff and rest of dorsum less ochraceous-olive; below, with the 
chin and fore-throat greyish white, rather than starkly white, and with 
the underside duller and less vibrant yellow. First-year dress with 
upper-parts duller and less yellowish ochraceous-buff than in adults; 
ventrally duller, less yellowish white. 
Wings: ¢¢ 47-52 (49.5), n=8; 92 49-51.2 (49.9) mm, n=9. 
Range. Western parts of the Namibian and Cape portions of the range 
from the Tropic of Capricorn south along the coast to the Cape of 
Good Hope and the Cape Agulhas region in the south, and east of A. m. 
damarensis to parts of southwestern Botswana in the northern Cape and 
the southern Transvaal at Hamanskraal. Generally occurs in areas with 
annual rainfall less than 120 mm. 


Anthoscopus minutus gigi Winterbottom, Bull. Brit. Orn. Cl. 79, p. 152. 
Oudtshoorn, southern Cape. 

Adult with upper-parts uniformly darker from the mantle to the rump 
(about dark Greyish Olive); the rump and upper tail-converts 
ochraceous-olive. Below, with the chin and fore-throat greyish or light 
fawn, tending to be darker laterally than in either of the other two forms; 
rest of venter olivaceous rather than yellow. Birds in first-year dress are 
ventrally lighter, with the lateral body surfaces and flanks washed with 
light brown rather than yellowish; dorsally they are a trifle darker. 

Wings: dod 48.5-52 (49.4), n=7; 99 49-52 (50.2) mm, n=11. 

Range. Southern and southeastern Cape from about 22°30'S, south of 
the mid-Orange and in the western parts of the Free State. Birds 
showing intergradation with minutus are present along the reaches of 
the lower Vaal R., near Kimberley. Generally confined to areas with 
annual rainfall of less than 250 mm. 


Acknowledgements 


For the loan of additional material to augment that available in Durban, I am indebted to 
the Ornithologists of both the East London and the National Museum, Bloemfontein. 


References: 

Clancey, P. A. 1980. S.A.O.S. Checklist of Southern African Birds. Southern African 
Ornithological Society, Johannesburg, Transvaal. 

Moreau, R. E. & Hall, B. P. 1970. An Atlas of Speciation in African Passerine Birds. 
British Museum (Nat. Hist.), London. 

Macdonald, J. D. 1952. The validity of Aegithalos smithi Jardine. Bull. Brit. Orn. Cl. 72: 
47-49. 

Ridgway, R. 1912. Color Standards & Color Nomenclature. The Author, Smithsonian 
Institution, Washington. 

Snow, D. W. 1967. Peters’ Checklist of Birds of the World. Vol. 12. Museum of 
Comparative Zoology, Harvard. 

Winterbottom, J. M. 1959. A new subspecies of Anthoscopus minutus (Shaw & Nodder) 
from Cape Province. Bull. Brit. Orn. Cl. 79: 152. 


Address: Dr P. A. Clancey, Suite 19, Caister Lodge, 264 Musgrave Road, Berea, Durban 
4001, South Africa. 


© British Ornithologists’ Club 1997 


C. J. Hazevoet 56 Bull. B.O.C. 1997 117(1) 


On a record of the Wattled Crane Bugeranus 
carunculatus from Guinea-Bissau 


by C. F. Hazevoet 
Received 28 December 1995 


The Wattled Crane Bugeranus carunculatus is a rare and declining 
Afrotropical species distributed in the Ethiopian highlands and in 
central and southern Africa, with a total population in 1983 of not more 
than 6000 and possibly as low as 4000 (Urban et al. 1986). Total 
population numbers given by Collar & Stuart (1985) are slightly more 
optimistic, viz. little more than 7500 and conceivably as low as 6000. 
There is a single record from West Africa of a party of three birds at 
Cufada lagoon, near Fulacunda, Guinea-Bissau (11°40’N, 14°55’W), 
supported by a photograph and a specimen (Frade & Bacelar 1955). 
The record was included (and considered “‘surprising’’) by Mackworth- 
Praed & Grant (1970), while Snow (1978), who did not map the record, 
remarked that “it seems almost certain that [the birds] were either 
escapees from captivity or vagrants’. Later, Collar & Stuart (1985) cast 
doubt on the validity of the record, stating that—referring to Frade 
& Bacelar (1955)—it was “‘supported without comment by a poor 
photograph which indicates that two or three birds were then present, 
given alongside an incomplete reference that suggests the species had 
previously been recorded at this locality’. The record was not 
mentioned by Urban et al. (1986), but recently the matter was raised 
again by Dowsett (1993) who stated that “the record appears 
astonishing from what is known of the distribution of the species, and 
even though it is supported by photos I think it best not to accept the 
record unconditionally, pending some further evidence’. Dowsett 
(1993) did not give the source of the record, nor did he mention that a 
specimen had been taken. 


The record 


The photograph was taken and the specimen collected by Major Julio 
de Araujo Ferreira, a Portuguese soldier and naturalist stationed at 
Bissau at the time, who spent five days observing and photographing 
wildlife in the Cufada region in late March 1948 (Ferreira 1948). His 
itinerary allows the precise date of the record to be determined, viz. 
26 March, the Friday before Easter Sunday. The photograph was 
reproduced in Frade & Bacelar (1955), but a better quality 
reproduction of the same picture can be found in Ferreira (1948), and 
this leaves no doubt whatsoever as to the identity of at least two of the 
three birds shown. The third bird is seen in back view, with its head 
away from the photographer, and does, therefore, perhaps not allow 
certain identification. Ferreira (1948), however, explicitly stated that he 
saw three birds and there appears to be no reason to doubt this. 
Apparently one of the birds was collected (but only partly preserved) 


C. ¥. Hazevoet 57 Bull. B.O.C. 1997 117(1) 


and a photograph of the skull was published in Frade & Bacelar (1955). 
It appears that, apart from the skull, only one leg was preserved and 
Frade & Bacelar (1955) gave the following measurements: bill 155 mm, 
tarsus 305 mm, middle toe 115 mm. They suggested that this bird was 
immature but this cannot be decided from the measurements or the 
photograph. The measurements are within the range given for adults (cf. 
Walkinshaw 1973, Maclean 1985, Urban et al. 1986). 

Although Ferreira’s (1948) paper is written in a popular fashion, it 
includes many original data, with scientific names employed 
throughout, and it was illustrated with several photographs of birds and 
mammals taken during his visit to the Cufada region. About the 
Wattled Cranes he remarked that they were accompanied by a 
Saddle-billed Stork Ephippiorhynchus senegalensis “‘which could be 
easily photographed from the canoe, but the cranes were very shy and 
this forced me to crawl through the marsh vegetation in order to 
approach them sufficiently close’ (translated). The conspicuous 
shyness of the species has also been reported by other authors, e.g. 
Collar & Stuart (1985) and references cited therein. 

Frade & Bacelar (1955, 1959) presented the ornithological results of 
the 1945-1946 Missado Zoologica to Portuguese Guinea, organized at the 
initiative of the Natural History Museum of Lisbon. Data were 
presented in a scholarly manner, giving details of all specimens 
collected during the expedition as well as full citations of all previous 
reports of species collected in the country. Collar & Stuart’s (1985) 
interpretation of Frade & Bacelar (1955), as given above, is less than 
adequate. In fact, the entry on B. carunculatus reads (translated): ‘‘Not 
collected during the Missdo, but a specimen collected by Major Aratyjo 
Ferreira was presented to us (head, leg and foot); No. 1-1948, Cufada 
lagoon, Fulacunda’’. They continued: “‘It appears that this species has, 
so far, not been recorded from West Africa north of Benguela (Angola) 
and its presence in Portuguese Guinea is indeed unexpected ... 
Bannerman does not mention it in any of his works on the birds of West 
Africa ... J. A. Ferreira, in his paper on the Fauna of the Cufada 
Reserve, mentioned the species under the indication grous de 
caruncula’. Contrary to Collar & Stuart’s (1985) interpretation, there is 
no suggestion whatsoever that the species had been recorded on any 
other occasion while the reference to Ferreira’s (1948) paper is given in 
full in the bibliography. Moreover, the caption of the photograph 
clearly stated that this was taken by ‘capitao Araujo Ferreira’. It 
appears that not all the authors who expressed their doubt about the 
record (i.e. Snow 1978, Collar & Stuart 1985, Dowsett 1993) had fully 
understood Frade & Bacelar’s (1955) Portuguese text, and that none of 
them checked the original paper by Ferreira (1948). 

The fate of the specimen is unknown at present. It was presented, 
together with other specimens collected by J. A. Ferreira, to the Missdo 
Zoologica (Frade & Bacelar 1955). The collections of the Missdo are 
kept at the Centro de Zoologia of the Instituto de Investiga¢ao 
Cientifica Tropical, Lisbon, Portugal, but the specimen of Bugeranus 
carunculatus could not be located there (M. M. Pinheiro in litt. 13 
August 1996). 


C. 7. Hazevoet 58 Bull. B.O.C. 1997 117(1) 


Discussion 


The Wattled Crane is a bird of local and discontinuous distribution and 
its movements are poorly understood. In Ethiopia it is present in 
highland marshes at the end of long rains but disappears as marshes dry 
up, and in southern Africa nomadic movements have been noted 
(Collar & Stuart 1985, Urban et al. 1986). Over the last few decades, 
the species has declined severely both in numbers and range and it 
now consists of two widely separated populations in Ethiopia and in 
central and southern Africa (Collar & Stuart 1985, Urban et al. 1986). 
This decline is attributed to loss of habitat, human interference 
and development projects, factors which undoubtedly play their role 
today. Initially, however, the separation of the Ethiopian and more 
southern populations may well have been due to natural fluctuations 
of the climate and subsequent changes in the hydrography and 
vegetation of the intervening areas, breaking up a once more or less 
continuous distribution (see e.g. Livingstone 1975, Zinderen Bakker 
1976). Snow (1978) presumed that the absence of any appreciable 
geographical variation indicated a good deal of movement between 
populations. Alternatively, the combination of discontinuous distri- 
bution and lack of geographical variation may be due to relatively 
recent vicariance events resulting from climatic change. Range 
fragmentation has presumably been accelerated by human activities in 
recent decades. 

In the past, the species may also have occurred further north, as it 
is depicted in Ancient Egyptian illustrations, indicating its former 
occurrence in lowland areas northwest of the Ethiopian highlands 
and perhaps even in Egypt itself (Kumerloeve 1983). During the 
Late Pleistocene, when at times a much more humid climate 
prevailed in the western part of the Sahara and the Sahel Zone than 
today (e.g. Nicholson & Flohn 1980, Dupont & Hooghiemstra 1989), 
the range of the Wattled Crane may have extended well into West 
Africa. One may speculate that the Guinea-Bissau birds were 
vagrants from Ethiopia, Zaire or Angola, having reached West Africa 
by way of the Lake Chad area, the Niger Inner Delta or other 
marshlands. On the other hand, there exists the possibility that the 
Guinea-Bissau birds were local breeders and belonged to a remnant 
and now extinct West African population. Although there is at 
present no further evidence for this, the possibility should at least be 
considered and not dismissed a priori. Indeed, the possibility of a 
fragmented range appears to be more parsimonious than the 
vagrant option, and this hypothesis would even gain in strength if 
one of the birds was indeed immature, as suggested by Frade & 
Bacelar (1955). 

Of course, the possibility of the Guinea-Bissau birds having been 
escapees, as suggested by Snow (1978), cannot be proven to be false but 
this appears highly unlikely taking into account the location and year of 
the record. Although the record is certainly ‘“‘surprising’’ (and perhaps 
even ‘‘astonishing’’), there is nothing to suggest that the birds were 
anything but genuinely wild. 


C. ¥. Hazevoet 59 Bull. B.O.C. 1997 117(1) 


Acknowledgements 


I wish to thank C. S. Roselaar and Jan Wattel for providing useful comments on the 
manuscript. Jan Wattel also worked out the date of Easter Sunday 1948. I thank Maria 
Margarida Pinheiro (Centro de Zoologia, Lisbon) for her efforts in trying to locate the 
specimen. 


References: 

Collar, N. J. & Stuart, S. N. 1985. Threatened Birds of Africa and Related Islands. 
International Council for Bird Preservation & International Union for Conservation 
of Nature and Natural Resources, Cambridge. 

Dowsett, R. J. 1993. Afrotropical avifaunas: annotated country checklists. Tauraco Res. 
Rep. 5: 1-322. 

Dupont, L. M. & Hooghiemstra, H. 1989. The Saharan-Sahelian boundary during the 
Brunhes chron. Acta Bot. Neerl. 38: 405-415. 

Ferreira, J. A. 1948. Fauna da reserva de Cufada. Bol. Cult. Guiné Port. 3: 739-758. 

Frade, F. & Bacelar, A. 1955. Catalogo das aves da Guiné portuguesa, I—Non-Passeres. 
Anais Junta Invest. Ultramar 10 (4, fasc. 2): 1-194. 

Frade, F. & Bacelar, A. 1959. Catalogo das aves da Guiné portuguesa, I]—Passeres. 
Mem. Junta Invest. Ultramar (Ser. 2) 7: 1-116. 

Kumerloeve, H. 1983. Zur Kenntnis altagyptischer Ibis-Darstellungen, unter besonderer 
Berticksichtigung des Waldrapps, Geronticus eremita (Linnaeus, 1758). Bonn. Zool. 
Beitr. 34: 197-234. 

Livingstone, D. A. 1975. Late Quaternary climatic change in Africa. Ann. Rev. Ecol. 
Syst. 6: 249-280. 

Mackworth-Praed, C. W. & Grant, C. H. B. 1970. Birds of West Central and Western 
Africa. African handbook of birds. Ser. 3, Vol. 1. Longmans, London. 

Maclean, G. L. 1985. Roberts’ Birds of Southern Africa. John Voelcker Bird Book Fund, 
Cape Town. 

Nicholson, S. & Flohn, H. 1980. African environmental and climatic changes and the 
general atmospheric circulation in Late Pleistocene and Holocene. Climatic Change 
2: 313-348. 

Snow, D. W. 1978. An Atlas of Speciation in African Non-passerine Birds. British 
Museum (Natural History), London. 

Urban, E. K., Fry, C. H. & Keith, S. 1986. The Birds of Africa. Vol. 2. Academic Press, 
London. 

Walkinshaw, L. H. 1973. Cranes of the World. Winchester Press, New York. 

Zinderen Bakker, E. M. van. 1976. The evolution of Late-Quaternary palaeoclimates of 
southern Africa. Palaéoecol. Afr. 9: 160-202. 


Address: C. J. Hazevoet, Institute of Systematics and Population Biology, University of 
Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands. 


© British Ornithologists’ Club 1997 


P. G. P. Ericson & L. A. Amarilla 60 Bull. B.O.C. 1997 117(1) 


First observations and new distributional data 
for birds in Paraguay 


by Per G. P. Encson & Luts Alberto Amarilla 
Received 26 Fanuary 1996 


The Paraguayan bird fauna has long been one of the least studied in 
South America. Beginning in the late 1970s, this situation has gradually 
changed, and rather rapidly so during the last ten years. With the 
publication by Hayes (1995), the status, distribution and biogeography 
of the Paraguayan avifauna has been evaluated and summarised in the 
light of available published and unpublished sources, and of extensive 
field work. Hayes’ book constitutes a welcome and solid foundation 
for further ornithological studies in Paraguay. The processes that 
determine the geographical and temporal distributions are still poorly 
understood, however, and additional important information is collected 
during almost every major field trip in the country. 

In this paper, new distributional data on several bird species are 
presented. The data derive from two different sources. The first is a 
large, unpublished collection of Paraguayan birds stored at the Swedish 
Museum of Natural History. During an 18-month collecting trip to 
Paraguay in 1946-47, Claés-Christian Olrog collected 1048 birds, and 
many other vertebrates, for the Swedish Museum of Natural History, 
mainly from the Chaco. 

The second is data collected during recent collaborative field work in 
Paraguay by the Museo Nacional de Historia Natural del Paraguay and 
the Swedish Museum of Natural History. Since 1993, the two 
museums have been collaborating in the project PROVEPA (Proyecto 
Vertebrados del Paraguay). The project contains a biodiversity training 
programme focusing on various aspects of vertebrate systematics and 
taxonomy, biological diversity estimation, and natural history museum 
practices. Within this framework, ornithological field work has been 
conducted in various areas of Paraguay. Information on the avifauna is 
regularly obtained in the process of training students in censusing 
methods, sound-recording and specimen collecting. Needless to say, all 
regular sight observations are also recorded. 

The biogeographical regions referred to below are those defined from 
vegetational and topographical features by Hayes (1995) (Fig. 1). 


Specimen data standards 


Measurements. Wing length—maximum length method, 1.e. with wing 
flattened against the ruler and the primaries straightened (cf. Svensson 
1993); tail length—from the root of the central pair of rectrices; 
culmen—distance from tip to the base of the foremost feathers of the 
forehead; bill depth—measured where the calipers rest comfortably or, 
on wedge-shaped bills, at the feathering with the calipers oriented at 
90 degrees against the cutting edge of the bill; tarsus length—from 


P. G. P. Ericson & L. A. Amarilla 61 Bull. B.O.C. 1997 117(1) 


AC Alto Chaco | 
MG Matogroséensé | 
BC Bajo Chaco tess; | 
| 
| 
| 


CC Campos Cerracos 
' CP Central Paraguay 


~ NE Neembucu 
| AP Alto Parana 


Figure 1. Biogeographical divisions of Paraguay based on vegetation and geographical 
features; from Hayes (1995). Alto Chaco, Matogrosense and Bajo Chaco constitute the 
Paraguayan Chaco. The region east of Rio Paraguay is called the Oriente. 


intertarsal joint to last complete leg scale before the toes diverge. All 
measurements are given in millimetres. 
Museum acronyms. NRM stands for the Swedish Museum of Natural 


History and MNHWNP for the Museo Nacional de Historia Natural del 
Paraguay. 


Annotated list of species 


RED-WINGED TINAMOU Rhynchotus rufescens pallescens 


The Red-winged Tinamou is widely distributed in Bolivia, Brazil, 
Paraguay, Uruguay and Argentina (Blake 1977). In Paraguay it is 
common only in the Oriente, and is regarded as not occurring in the 
Alto Chaco and Matogrosense biogeographical regions. 

Three males and five females (NRM 500061-500068), collected by 
Olrog in western Dpto. Presidente Hayes in the austral winter of 1947, 
constitute the first documented records of the Red-winged Tinamou 


P.G.P. Ericson & L.A. Amarilla 62 Bull. B.O.C. 1997 117(1) 


TABLE 1 
Size comparisons (mm) of two subspecies of the Red-winged Tinamou Rhynchotus 
rufescens. Comparative data from Blake (1977) 


Wing length Culmen 
Males 
R. r. rufescens 10 192-207 36-39 
R. r. pallescens 5 211-229 36—42 
Olrog coll. 3 196-215 40-42 
Females 
R. r. rufescens 10 183-211 34-39 
R. r. pallescens 10 205-226 36-42 
Olrog coll. 5 195-217 36-44 


for the Alto Chaco biogeographical region. The birds collected by 
Olrog are much greyer on the breast and the belly than the nominate 
subspecies, to which the Chaco population has been assigned (Short 
1975). Instead, they fully match the coloration of the race R. r. 
pallescens, that occurs in the grassy lowlands of north-central 
Argentina. From the rather small Paraguayan sample, it seems that the 
Chaco population of R. r. pallescens is somewhat smaller than are 
typical R. r. pallescens (Table 1). The distribution of this subspecies in 
the Paraguayan Chaco, in relation to that of the nominate subspecies 
(cf. Short 1975), remains to be determined. 


HOOK-BILLED KITE Chondrohierax uncinatus uncinatus 

The Hook-billed Kite occurs widely in Middle and South America. 
In Paraguay it is a rare resident in several biogeographical regions 
in both the Chaco and Oriente. A female (NRM 500111) collected 
23 November 1946 c. 40 km east of Pozo Colorado in Dpto. Presidente 
Hayes is, however, the first record of the species from the Bajo Chaco 
region. The active egg follicles show that the bird was breeding at the 
time. 


BLACK-BANDED OWL Ciccaba huhula 


The Black-banded Owl has a wide distribution in forests of northern 
and eastern South America and occurs nearest to Paraguay in eastern 
Brazil (Meyer de Schauensee 1966). The only previous report of the 
Black-banded Owl from Paraguay is based on calls attributed to this 
species at the Estancia Itab6, Dpto. Canindeyu, by Brooks e¢ al. (1992). 

On 30 September 1993 a female Black-banded Owl (NRM field 
no. ICM-006; gonads active) was collected in Parque Nacional 
Caaguazu, Dpto. Caazapé, by a PROVEPA team. The owl was 
breeding, as shown by an almost fully developed egg present in the 
ovary with additional eggs under development. Two bats of different 
species were found in the stomach. Nothing is known of the food 
preferences of the Black-banded Owl, but information does exist on the 


P.G. P. Ericson & L.A. Amarilla 63 Bull. B.O.C. 1997 117(1) 


diet of its allospecies, the Black-and-white Owl Ciccaba nigrolineata. 
That species frequently captures bats (Ibanez et al. 1992), and other 
data suggest that it feeds mainly on large insects (Mikkola 1973). 

Despite considerable ornithological activity in areas where the 
Black-banded Owl could be expected to occur, it has never been 
recorded with certainty before (contra Meyer de Schauensee 1966). 
This makes it likely that the Paraguayan population of the species is 
rather small and local, although this remains to be determined. If so, 
the Black-banded Owl certainly warrants protection in Paraguay. 


BLUE-CROWNED MOTMOT Momotus momota 

Although widely distributed in tropical South America, the 
Blue-crowned Motmot has only been sighted twice before in Paraguay, 
both times in the northeastern Chaco (Hayes et al. 1990). A male 
(MNHNP field no. ICM-078) obtained on 12 November 1994 by 
PROVEPA near the city Bella Vista in Dpto. Amambay, one of two 
birds observed to cross the Rio Apa from Brazil, is the first record from 
the Oriente. Two additional specimens, one unsexed and one male 
(NRM field nos. GFK-193 and PMN-145), collected 26-27 March 
1995 at the Parque Nacional Defensores del Chaco constitute the first 
from the Alto Chaco biogeographical region. 

From their distribution, these specimens are most likely to belong to 
the subspecies M. m. pilcomajensis, which occurs in southern Bolivia, 
northern Argentina, and eastward to Urucum and possibly western S4o 
Paulo in Brazil (Chapman 1923). This subspecies is distinguished by 
its bluish-green throat and greenish underparts. The Paraguayan 
specimens are however much less bluish-green on throat than typical 
M. m. pilcomajensis from Argentina in the NRM collection, thus 
approaching the more rufous condition of some NRM specimens of the 
subspecies M. m. simplex from Amazonian Brazil. Also the breast and 
underparts are more cinnamon than typical M. m. pilcomajensis. It thus 
seems probable that these specimens are intergrades between M. m. 
pilcomajensis and simplex. 


RUSTY-BACKED ANTWREN Formicivora rufa rufa 


The Rusty-backed Antwren has a wide distribution in central South 
America (Ridgely & Tudor 1994). In Paraguay, it is a rare and local 
breeder in northern and central Oriente (Hayes 1995). Only one record 
exists from the Chaco: Villa Hayes in Dpto. Presidente Hayes (Bertoni 
1930, Hayes 1995). Its closest breeding areas outside Paraguay are in 
northeastern Bolivia and southern central Brazil (Meyer de Schauensee 
1966). 

On 16 September 1994, an adult male (NRM field no. GFK-111; tail 
moulting, gonads inactive) was obtained by a PROVEPA team 5 km 
north of Bahia Negra, Dpto. Alto Paraguay. This specimen is the 
first documented occurrence of the Rusty-backed Antwren in the 
Matogrosense biogeographical region. It was mist-netted in open forest 
with high scrub vegetation, only a couple of metres from the Rio 
Paraguay, indicating that it might have been a stray individual from the 
Mato Grosso do Sul of Brazil, where it is known to breed. 


P.G. P. Enicson & L. A. Amarilla 64 Bull. B.O.C. 1997 117(1) 


BRAN-COLOURED FLYCATCHER Myiophobus fasciatus 

The Bran-coloured Flycatcher has a wide, almost circum-Amazonian 
distribution in South America. Contrary to the map in Ridgely & 
Tudor (1994), this species has previously not been thoroughly 
documented from the Paraguayan Chaco (Hayes 1995). A juvenile 
(NRM field no. GFK-184; fresh plumage) obtained on 25 March 1995 
by a PROVEPA team in Parque Nacienal Defensores del Chaco, Dpto. 
Alto Chaco, thus constitutes the first Paraguayan record of the 
Bran-coloured Flycatcher west of the Rio Paraguay. 

The subspecific allocation of this individual is somewhat problem- 
atic. The subspecies mainly differ in size and in the amount of rufous in 
the dorsal coloration. In the collections of NRM we compared the 
PROVEPA bird with samples of M. f. auriceps and M. f. flammiceps 
taken in Bolivia, Argentina and Brazil. Although a considerable 
variation in dorsal coloration and size can be observed between the 
samples, this does not co-vary geographically. Instead, much of this 
variation seems to be due to differences in the age composition of the 
samples, the juveniles being considerably more reddish on the dorsal 
parts than the adults. It is suggested from the NRM collection that the 
southern populations of the Bran-coloured Flycatcher are in need of a 
taxonomic revision. 


RUSSET-WINGED SPADEBILL Platyrinchus leucoryphus 

The Russet-winged Spadebill is endemic to the Mata Atlantica with 
a very restricted distribution in southeastern Brazil, eastern Paraguay 
and northeastern Argentina. It is rare and regarded as Vulnerable by 
Collar et al. (1992), who stressed the need for further field studies of its 
population dynamics and habitat requirements, which are largely 
unknown. 

On 6 October 1993 an adult male (NRM field no. ICM-020) 
Russet-winged Spadebill was mist-netted by a PROVEPA team in 
primary forest at Parque Nacional Caaguazi in Dpto. Caazapa, 
Paraguay. Its testes were enlarged, indicating breeding condition. The 
stomach contained invertebrates in the following proportions by 
weight: 15% Hymenoptera, 5% Araneae, 5% Coleoptera, 4% Diptera? 
(eggs), 1% Arthropoda (eggs), and 70% undetermined. At the same site 
several individuals of the smaller, congeneric White-throated Spadebill 
P. mystaceus were obtained. Comparisons of the stomach contents of 
these few individuals could reveal no significant differences in food 
choice between the species. 

The Russet-winged Spadebill is a rather inconspicuous bird 
occurring in the understory of subtropical forests. Albeit never in large 
numbers, the species has been collected and sighted at several localities 
in the Oriente where it may be more common than previously thought. 


STRANGE-TAILED TYRANT Alectrurus risora 

The Strange-tailed Tyrant is declining or has become extinct in large 
areas of its former distribution in southern Brazil, Paraguay, Uruguay 
and northern Argentina, mainly due to changes in land management 
(Collar et al. 1992). The present Paraguayan distribution is mainly in 


P. G. P. Ericson & L. A. Amarilla 65 Bull. B.O.C. 1997 117(1) 


the southernmost part of the Oriente and the humid Chaco where it 
is local and uncommon (Collar et al. 1992, Hayes 1995). Two males 
and one female (MNHNP field no. AHN-203, and NRM field 
nos. AHN-206 and AHN-201; no moult recorded and gonads inactive) 
were obtained on 28 September 1994 at Estancia San José, Dpto. 
Presidente Hayes, where it is a not uncommon breeder (pers. obs.). 


WHITE-NAPED XENOPSARIS Xenopsaris albinucha albinucha 


In areas bordering Paraguay, the White-naped Xenopsaris occurs in 
northern Argentina, eastern Bolivia and southwestern Brazil, where 
it seems to be mostly a rare and local species (Ridgely & Tudor 1994). 
On 22 November 1946, Olrog collected a male (NRM_ 500445) 
White-naped Xenopsaris at Estancia Hermosa, c. 40 km east of Pozo 
Colorado, Dpto. Presidente Hayes. This is the first record of the 
species from the Bajo Chaco, despite a good number of records from 
the Alto Chaco biogeographical region of Paraguay (Hayes 1995). 


GOLDEN-BILLED SALTATOR Saltator aurantitrostris aurantitrostris 


Although the Golden-billed Saltator is generally distributed in 
western Paraguay (Ridgely & ‘Tudor 1989) and is even abundant in the 
Alto Chaco biogeographical region, it has never been reported from the 
Matogrosense (Hayes 1995). On 13 September 1994, a PROVEPA 
team obtained a female (NRM field no. NEY-038; not moulting) at 
Estancia Dona Julia, 5 km north of Bahia Negra, Dpto. Alto Paraguay. 


ULTRAMARINE GROSBEAK Cyanocompsa brissonii sterea 


The Ultramarine Grosbeak is a rare to uncommon breeder in most 
areas of Paraguay (Hayes 1995). A specimen (NRM field no. LIY-019) 
obtained 13 October 1994 by a PROVEPA team near the city of Bella 
Vista, Dpto. Amambay, is the first record from the Campos Cerrados 
biogeographical region. Its small size (wing 80, tail 75, culmen 14.5, bill 
depth 13, tarsus 19) shows that it should be referred to the subspecies 
C. b. sterea, of eastern and southern Brazil, northeastern Argentina, and 
western Paraguay (Oberholser 1901, Paynter 1970). 


SAFFRON-COWLED BLACKBIRD A gelaius flavus 


The Saffron-cowled Blackbird is a rare breeder in the Oriente region 
of Paraguay, from which only a handful of reported sightings exist 
(Hayes 1995). On 3 March 1995, Per Ericson and Ingrid Cederholm 
observed four male Saffron-cowled Blackbirds perched for 2 minutes in 
a tree along Ruta General Bruguez, Dpto. Presidente Hayes (c. 24°45'S, 
58°30'W). The typical icterid appearance of these birds, in combination 
with a yellow head, leaves no doubt about the species identification. 
This observation, made in open, wet grassland, is the first documented 
sighting in the Chaco of Paraguay. The species may, however, have 
been observed at least once before in the southern Bajo Chaco (Collar 
et al. 1992, Jorge Escobar pers. comm.). Because of its declining 
populations in all parts of its range, the Saffron-cowled Blackbird is 
regarded as Vulnerable (Collar et al. 1992). 


P.G. P. Ericson & L. A. Amarilla 66 Bull. B.O.C. 1997 117(1) 


TROUPIAL Icterus icterus strictifrons 


In Paraguay, the 'Troupial occurs in the Alto Chaco and Bajo Chaco 
biogeographical regions. On 17 September 1994 an adult male (NRM 
field no. GFK-130; gonads active) was collected in the Matogrosense 
region by a PROVEPA team 5 km north of Bahia Negra, Dpto. Alto 
Paraguay. The bird appeared to be paired with another individual, but 
apart from its active gonads there was no further sign of breeding 
activity. 

The two Troupials were the only individuals of the species observed 
during the 12 days of field work in the area. Since the collecting site is 
only 20 m from the Rio Paraguay, the birds may have belonged to the 
population that inhabits the Rio Paraguay drainage of the Brazilian 
Mato Grosso (Ridgely & Tudor 1989). 


Acknowledgements 


We thank Ingrid Cederholm, Jorge Escobar, Nubia Etcheverry, Rosalia Farina, Goran 
Frisk, Anders Hansson, Lars Imby, Cristina Morales and Peter Mortensen, who 
participated as ornithologists and scientific preparators in the PROVEPA project. The 
Direccion de Parques Nacionales y Vida Silvestre, Asuncion, and the Museo Nacional de 
Historia Natural, San Lorenzo, provided PROVEPA with logistical service and 
permitted work on government property. In addition, we thank the private landowners 
who let us work on their properties. We offer special thanks also to Tomas Johansson, 
who did the painstaking work of checking all the specimen labels in the Olrog collection 
against the computerised card file at the Swedish Museum of Natural History. The 
determinations of the invertebrates found in the gizzards were coordinated by Stefan 
Lundgren, Service Centre for Taxonomic Zoology, Swedish Museum of Natural 
History. Thanks also to Drs Robert S. Ridgely, J. Van Remsen, Sven O. Kullander and 
Goran Frisk for commenting on earlier drafts of the manuscript. 


References: 

Bertoni, A. de W. 1930. Sobre ornitologia del Chaco Paraguayo. Aves colectadas por 
Félix Posner en la Colonia ‘‘Monte Sociedad’’, hoy Benjamin Aceval (Villa Hayes). 
Rev. Soc. Cient. Parag. 2: 241-257. 

Blake, E. R. 1977. Manual of Neotropical Birds. Univ. Chicago Press. 

Brooks, T. M., Barnes, R., Bartrina, L., Butchart, S. H. M., Clay, R. P., Esquivel, E. Z., 
Etcheverry, N. I., Lowen, J. C. & Vincent, J. 1992. Bird surveys and conservation in 
the Paraguayan Atlantic forest. BirdLife Int. Study Rep. 57. 

Chapman, F. M. 1923. The distribution of the motmots of the genus Momotus. Bull. Am. 
Mus. Nat. Hist. 48: 27-59. 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
T. A. & Wege, D. C. 1992. Threatened Birds of the Americas. The ICBP/IUCN Red 
Data Book. Smithsonian Institution Press. 

Hayes, F. E. 1995. Status, distribution and biogeography of the birds of Paraguay. 
Monogr. Field Ornithol. 1. Colorado Springs: American Birding Association. 

Hayes, F., Goodman, S. M. & Lopez, N. E. 1990. New and noteworthy bird records 
from the Matogrosense region of Paraguay. Bull. Brit. Orn. Cl. 110: 94-103. 

Ibanez, C., Ramo, C. & Busto, B. 1992. Notes on food habits of the Black-and-white 
Owl. Condor 94: 529-531. 

Meyer de Schauensee, R. 1966. Birds of South America. Academy of Natural Sciences of 
Philadelphia. 

Mikkola, H. 1973. Wood owls. Jn: J. A. Burton (ed.), Owls of the World, their evolution, 
structure and ecology. Peter Lowe, Amsterdam. 

Oberholser, H. C. 1901. Seven new birds from Paraguay. Proc. Biol. Soc. Washington 14: 
187-188. 

Paynter, R. A. Jr. 1970. Subfamily Cardinalinae. Im: R. A. Paynter, Jr. (ed.), Check-list of 
the Birds of the World. Vol. 13. Museum of Comparative Zoology, Harvard. 


G. F! Mees 67 Bull. B.O.C. 1997 117(1) 


Ridgely, R. S. & Tudor, G. 1989. The Birds of South America. Vol. 1. The Oscine 
Passerines. Univ. of Texas Press. 

Ridgely, R. S. & Tudor, G. 1994. The Birds of South America. Vol. 2. The Suboscine 
Passerines. Oxford Univ. Press. 

Short, L. L. 1975. A zoogeographic analysis of the South American Chaco fauna. Bull. 
Am. Mus. Nat. Hist. 154: 1-82. 

Svensson, L. 1993. Identification Guide to European Passerines. Stockholm. 


Addresses: P. G. P. Ericson, Swedish Museum of Natural History, P.O. Box 50007, 
S-104 05 Stockholm, Sweden. L. A. Amarilla, Museo Nacional de Historia Natural 
del] Paraguay, Sucursal 19, San Lorenzo Km 9.5, Paraguay. 


© British Ornithologists’ Club 1997 


On the identity of Heterornis senex Bonaparte 


by G. F. Mees 


Received 10 January 1996 


The name Heterornis senex was introduced in the ornithological 
literature by Bonaparte (1850: 419) with the following words: ‘Pastor 
senex, Temm. Mus. Lugd. ex Beng. Dorso brunnescente, alis caudaque 
chalybaeo-nigris, pileo griseo; subtus sordide albo-cinereus”’. ‘The 
citation shows that, in Leiden, the mounted bird was labelled with the 
unpublished name Pastor senex 'Temminck. The binomen under which 
Bonaparte published the name was Heterornis senex. 

Only a few years later, Layard (1854: 217) described a mountain- 
starling from Ceylon, which he named Heterornis albofrontata. In the 
discussion, he noted: “‘It may be Pastor Senex, 'Temm., as it agrees 
tolerably well with the short description given in Prince Bonaparte’s 
Consp. Avium, p. 419, but that description is so concise that I cannot 
be sure of it; I therefore name it provisionally H. albofrontata’’. 

Layard’s suggestion that H. senex and H. albofrontata could be 
identical was soon transformed into certainty by Holdsworth (1872; 
462): ““Temenuchus senex, 'Temm. Peculiar to Ceylon; described by 
Layard as T. albofrontatus, as it was believed to be new; it has since 
been recognized as TJ. senex, "Temm., erroneously described by 
Bonaparte from Bengal’’. 

Since that time, the Ceylonese mountain-starling has been 
universally known by the specific epithet senex (its generic name has 
been less stable, the current one is Sturnus). 

In spite of Holdsworth’s definite statement, I have been unable to 
find published evidence that, since Bonaparte, anybody has ever 
actually examined the type material of H. senex. 

In the old collection of mounted birds in Leiden, there are two 
specimens labelled Pastor senex. Both have a red-margined label for 
display, of a type used in Temminck’s day (i.e. before 1858). The first 


G. F. Mees 68 Bull. B.O.C. 1997 117(1) 


one is labelled: ‘Pastor senex/'Temm. n. sp./ Bp. consp. / jeune / 
Chine’’, and underneath the socle only a number: 45. This bird has 
evidently been provided with a new pedestal at some date after it was 
examined by Bonaparte, as it was ‘Temminck’s habit to write the 
name and other particulars underneath the socles of mounted 
specimens. The other bird has, underneath the socle, in TTemminck’s 
handwriting: ‘Pastor senex § ad./'Temm. Nov. Sp./ Bp. consp. / 
Chine’’, and on the red-margined label exactly the same (the hand- 
writing on the red-margined labels is unfamiliar, probably by a clerical 
assistant). 

‘These birds have nothing to do with the Ceylonese species at present 
bearing their name, but are Sturnus sericeus Gmelin, 1789. The adult 
bird has a black pileum, and its identification requires no comment. 
The juvenile bird is a specimen of considerable antiquity; the way it is 
mounted suggests that it dates from the late 18th or early 19th century. 
Its upper parts are tinged with brownish and it lacks the black cap of 
the adult. As to their type status, Bonaparte’s description was clearly 
based only on the juvenile specimen, which therefore is the holotype. 

Regarding the authenticity of these two specimens, it certainly is a 
pity that the socle of the juvenile bird was replaced. It should be noted 
further that the provenance Bengal, given by Bonaparte, has been 
changed to China. The reference to Bonaparte’s Conspectus also means 
that the labels could not have been written before 1850. Perhaps the 
second, adult, specimen was received from China only a short time 
after Bonaparte examined the juvenile bird, and ‘Temminck may have 
changed the provenance of the latter to agree with this new 
information. Alternatively, the locality Bengal given by Bonaparte may 
simply be an error, of a kind numerous in his work. 

Although I have been aware of the true identity of these birds for 
some years, it should be stressed that otherwise they seem never to have 
been examined or studied since c. 1850. These two specimens are still 
the only Sturnus sericeus in the Leiden collection, and the Ceylonese 
species is even now unrepresented in Leiden. This probably explains 
why none of my predecessors in Leiden ever examined these birds. 

The question of nomenclature of the Ceylonese species remains. 
Evidently and simply, its correct name is Sturnus albofrontatus (Layard, 


1854). 


References: 

Bonaparte, C. L. 1850. Conspectus Generum Avium. Vol. 1. Lugduno Batavorum. 

Holdsworth, E. W. H. 1872. Catalogue of the birds of Ceylon; with some remarks on 
their habits and local distribution, and descriptions of two new species peculiar to 
the island. Proc. Zool. Soc. London: 404-483, pl. XVII-XX. 

Layard, E. L. 1854. Notes on the ornithology of Ceylon, collected during an eight years’ 
residence in the island. Ann. Mag. Nat. Hist. (2) 13: 212-218 (nos. 146-157). 


Address: G. F. Mees, 31 West Street, Busselton 6280, Western Australia. 


© British Ornithologists’ Club 1997 


Fearne al. 69 Bull. B.O.C. 1997 117(1) 


New distributional information on eight bird 
species from northern Peru 


by Roger Barnes, Stuart H. M. Butchart, 
Charles W. N. Davies, Mirko Fernandez & 
Nathalie Seddon 


Received 21 March 1996 


In July and August 1994 our team of five British and Peruvian 
ornithologists carried out surveys during an expedition organized with 
the help of the University of Cambridge, in close consultation with the 
Asociacion Peruana para la Conservacion de la Naturaleza (APECO). 
Our principle aim was to assess the conservation status and habitat 
requirements of threatened and endemic birds and mammals in 
remaining areas of cloudforest and elfin forest on the Cordillera de 
Colan in Amazonas Department, northern Peru. 

The Cordillera de Colan is a large, projecting mountain ridge with 
high rainfall. Its forest cover is a complex mosaic of habitats, some of 
only limited extent. Human activity has altered and degraded several of 
these habitats (Davies et al. 1994). The range is almost surrounded by 
the dry forest in the valleys of the rio Maranon and its tributaries, 
which holds its own endemic avifauna. 

We concentrated our fieldwork at altitudes of 1500—2650 m at two 
sites in the southern part of the range, with some observations down 
to 550m (Fig. 1, Table 1). Amongst our records were extensions of 
the known distribution of eight bird species. One of them, the Royal 
Sunangel Heliangelus regalis, is the subject of a previously published 
paper (Seddon et al. 1996). 


WHITE-EYED PARAKEET A ratinga leucophthalmus 


A widespread and fairly common species east of the Andes, recorded 
up to 500 m asl (Hilty & Brown 1986). We found it to be common at 
site 2, in flocks of up to 47, recording it daily from 17 to 30 August at 
700-1950 m in both cloudforest and humid lower montane elfin forest, 
where it was regularly seen at the ridgetop at 1950 m. 


WHITE-TIPPED SICKLEBILL Eutoxeres aquila 


A bird of lower montane humid forest, previously known from Costa 
Rica to Amazonas north of rio Maranon, northwest Peru (Hilty & 
Brown 1986). One was seen on 29 July at 2300 m in very humid lower 
montane cloudforest at site 1; we then netted and photographed four at 
1950m in ridgetop elfin forest at site 2 on 20-23 August. The 
Cordillera de Colan is a new site for this hummingbird and the first 
south of the deep gap formed by the Maranon valley which was 
previously presumed to restrict the range of this species (J. Fjeldsa in 
litt.). 


R. Barnes et al. 


Ecuador 


70 Bull. B.O.C. 1997 117(1) 


Amazonas 


Amazonas 


Pacific 
Ocean 


Loreto 


ys 
% 
O 


oo 
Cordillera: , 
“de Colan~ 


Zw 
ite 2: 


Bagua 
Grande 


“wShe 1: 
Comboca 


San Cristobal 


Rio Utcubamba 


Cajamarca 


San Martin 


Figure 1. The location of our study sites on the Cordillera de Colan, northern Peru. 


R. Barnes et al. 71 Bull. B.O.C. 1997 117(1) 


TABLE 1 
Site characteristics and fieldwork effort 


Altitudes Field-hours. Mistnet-hours 
Fieldwork surveyed ee ee Eee 
Site dates (m) Habitat day night day night 
1 13-15 Jul 550-1500 Humid premontane Casual obs. 
forest 
15-22 Jul 1500-2150 Humid lower 219 $5 129 
montane forest 
22 Jul-6 Aug 2150-2650 Very humid lower 286 8 3t9 DS) 
montane forest 
2 12-16 Aug 900-1100 Dry premontane Casual obs. 
forest 
15-31 Aug 1650-1850 Humid lower 149 3 
montane cloud 
forest 
15-31 Aug 1650-1750+ Humid lower 97 14 334 54 


1850-1950 montane elfin forest 


RUSSET-MANTLED SOFTTAIL Thripophaga berlepschi 


A very local species confined to the Andes of north Peru (Amazonas 
on Cordillera de Colan south to east Libertad) at 2450-3350 m (Ridgely 
& Tudor 1994). It is considered to be ‘“‘near-threatened”’ (Collar et al. 
1994). Restricted-range in EBA B25: North-east Peruvian cordilleras 
(Stattersfield et al. 1995). 

We had six records at 1800-1950 m, on 15-28 August, at site 2, all in 
ridgetop elfin forest except for one in adjacent humid lower montane 
cloudforest. The birds were most often seen foraging in the canopy 
with mixed-species flocks including Grey-mantled Wren Odontorchilus 
branicku. These records constitute a downward altitudinal range 
extension of 650 m for the species. 


BARRED ANTTHRUSH Chamaeza mollissima 


Rare and local in the Andes of Colombia, Ecuador and northern Peru 
(Cerro Chinguela in Piura), also the Andes of southern Peru (Cuzco 
and Puno) and western Bolivia. Mostly at 1800 to 3000 m asl. Ridgely 
& Tudor (1994) suggested that it may occur in the intervening section 
of the Andes of Peru. We recorded nine at 2300-2600 m in very humid 
upper montane forest at site 1, 24 July to 2 August, and one was netted 
and photographed at 2300 m on 26 July. 

These records occur between the two disjunct areas previously known 
for this “‘superb, rare antthrush” (Ridgely & Tudor 1994). The birds 
most closely resembled the northern nominate race mollissima, showing a 
barred rather than streaked throat, and narrow bars on the belly rather 
than coarse wavy barring. However the breast was not as clearly barred 
as on mollissima, and the markings suggested the chevroned breast 
pattern of the southern yungae (Ridgely & Tudor 1994). The Cordillera 


R. Barnes et al. 72 Bull. B.O.C. 1997 117(1) 


de Colan is much closer to the known range of mollissima than that of 
yungae, so the resemblance to the northern form is not surprising; but 
our records suggest plumage intermediacy in the intervening parts of 
this species’ range, and that the rio Maranon does not form a clean break 
between the two subspecies as might have been expected. 


CHESTNUT ANTPITTA Grallaria blakei 


A “‘near-threatened”’ species (Collar et al. 1994), with a restricted 
range in EBA B25: Northeast Peruvian cordilleras (Stattersfield et al. 
1995). Ridgely & ‘Tudor (1994) describe it as local on the eastern slope 
of the Andes in northern and central Peru in Amazonas, Huanuco and 
Pasco; 2150-2475 m. We recorded it once, on 16 July 1994 at 1700 m in 
humid lower montane forest at site 1. This small antpitta showed a 
black bill, greyish legs and uniform chestnut plumage with only the 
faintest hint of barring on the lower belly and no discernible eye ring. 
The bird was flushed from dense undergrowth, perched at 1m and 
called once: a brief weeoo. No other antpitta exhibits this combination 
of field characters. 

This record represents a 450 m downward extension of this species’ 
known altitudinal range. Although recorded only once during the 
project it was familiar to local people and said to be occasionally seen on 
forest tracks. 


RUSTY-TINGED ANTPITTA Grallaria przewalskiu 


This antpitta has a restricted range in EBA B25: North-east Peruvian 
cordilleras (Stattersfield et al. 1995). It has been recorded at 
2200-2750 m (Ridgely & Tudor 1994). 

This species was recorded 261 times, mainly aurally, at site 1 at 
1700-2650 m on 16 July to 9 August 1994; it was uncommon in humid 
lower montane cloudforest below 2150 m, but more common. in very 
humid lower montane cloudforest above this altitude. These records 
constitute a 500 m downward extension of the altitudinal range of this 
still poorly known species. 


BLACK-CHESTED FRUITEATER Pipreola lubomirskiu 


This species is local on the east slope of the Andes from southern 
Colombia to northern Peru (west of the rio Maranon in Cajamarca); 
1500-2300 m (Ridgely & Tudor 1994). It is considered to be “near- 
threatened’ by Collar et al. (1994). We had a total of six records, all in 
humid lower montane forest. It was rare at site 1, with two sightings on 
16 July at 1750 m, possibly of the same individual. It was uncommon at 
site 2: two birds were netted in elfin forest at 1900 m (a female on 21 
August and a male on 22 August), and two were seen, one in 
cloudforest at 1800 m on 29 August and one in elfin forest at 1900 m on 
26 August. A photograph of the male was published in Cotinga 2: 40. 

Our records clearly fit this species, with the males showing a 
distinctive solid black bib, the females a solid green bib, and both sexes 
lacking the pale tertial tips of the much commoner Green-and-black 
Fruiteater P. riefferiiz. The description precludes the closely related 


R. Barnes et al. 73 Bull. B.O.C. 1997 117(1) 


Masked Fruiteater P. pulchra, in which the male has an orange lower 
throat and upper breast, and the female is entirely green streaked 
yellow below. Previously, pulchra was thought to replace lubomirski 
south and east of the rio Maranon in Amazonas department southwards 
(Snow 1982, Ridgely & Tudor 1994), which would include the 
Cordillera de Colan. No pulchra were recorded on the mountain range. 

These records show that the rio Maranon does not clearly separate 
the ranges of lubomirskii and pulchra and suggest the possibility that 
they may be sympatric. Further fieldwork is obviously needed at other 
sites in Amazonas department. 


RED-BILLED TYRANNULET Zimmerius cinereicapillus 

This species has a restricted range in EBA B29: East Andean foothills 
(Stattersfield et al. 1995). It occurs locally on the eastern slope of the 
Andes in eastern Ecuador (known definitely only from western Napo) 
and eastern Peru (Huanuco to eastern Cuzco and Madre de Dios); 
750-1200 m. It was thought to possibly also occur in the intervening 
part of the Andes of Peru by Ridgely & Tudor (1994). 

It is not easy to identify, and was recorded with certainty only once: 
at site 2 at 1800 m in ridgetop elfin forest, on 30 August. The following 
field characters were noted: a mainly yellow and olive bird with the 
contrasting remiges typical of Zimmerius tyrannulets. The rounded 
head showed a soft grey cap blending to olive on the nape. An indistinct 
pale supercilium, broken pale eye ring, pale iris and dark eye stripe 
leading to olive-grey ear coverts gave an overall plain facial appearance 
ruling out Golden-faced Tyrannulet Z. chrysops. 'The short bill was 
blackish above but dark pink below, paler at the base. The upperparts 
were uniform olive green from mantle to tail, and the rectrices darker 
with pale fringing. All the remiges appeared dark olive with strong 
yellow fringes. The dirty-white throat and faintly streaked upper breast 
blended into the buffy yellow belly and under tail coverts. The bird 
called often, either a single seep or a trill with a distinct szp szp ending. 

This is the first sighting within the range suggested as likely by 
Ridgely & Tudor (1994). It also constitutes an upward altitudinal range 
extension of 600 m, though this may not be significant as the bird was 
with a small flock crossing through ridgetop elfin forest between valleys. 
The bird’s occurrence in such low canopy made identification easier, and 
the species may have been overlooked elsewhere on the Cordillera. 


BLACK-CHESTED MOUNTAIN TANAGER Buthraupis eximia 


This is an uncommon and local species ranging from the Andes of 
southwestern Venezuela to extreme northern Peru (Cerro Chinguela on 
Piura—Cajamarca border), mostly at 2800-3500 m (Ridgely & ‘Tudor 
1994). It inhabits elfin forest, low mossy old forest, humid shrubbery and 
Polylepis forest (Fjeldsa & Krabbe 1990). Two were seen at site 1 on 2 
August and one on 3 August, at 2500 m in very humid lower montane 
forest. The combination of blue crown and nape, green back and wings and 
black upper breast distinguishes it from the other mountain tanagers with 
mostly yellow underparts. Our records are the first south of the Maranon 
Valley for this rather inconspicuous species (Hilty & Brown 1986). 


R. Barnes et al. 74 Bull. B.O.C. 1997 117(1) 


Conclusions 

Our records of White-tipped Sicklebill, Black-chested Fruiteater and 
Black-chested Mountain Tanager, showing new range extensions across 
the Maranon gap, tend to corroborate Fjeldsa’s contention that this 
barrier is not as biogeographically significant as previously assumed 
(Fjeldsa 1994). There are at least nine other species with restricted 
ranges which are found north and south of the valley (Davies et al. 1994). 

Our new altitudinal records probably result from several factors, 
including the atypical habitat spread. The areas of elfin forest at unusually 
low altitude ‘“‘may be explained by a cool local climate owing to frequent 
rainfall’ (Collar et al. 1994). Thus local conditions possibly lead to some 
vegetational zones occurring at lower altitudes than usual, in turn perhaps 
explaining the records of Russet-mantled Softtail, Chestnut Antpitta and 
Rusty-tinged Antpitta at unusually low altitudes. 

The fascinating Cordillera de Colan is a biogeographically very com- 
plex mountain range, which supports taxa representative of six different 
EBAs (Davies et al. 1994). This vulnerable site remains in need of 
further study and, more importantly, needs urgent, effective protection. 


Acknowledgements 


The advisors and sponsors of the expedition during which these records were made are 
acknowledged in our report (Davies et al. 1994). We would also like to particularly thank 
our patron, the late Gerald Durrell; Dr Mariella Leo Luna and the Asociacion Peruana 
para la Conservacion de la Naturaleza; Thomas Schulenburg; Rob Clay; Jon Fjeldsa; staff 
at BirdLife International, Cambridge, especially David Wege; Susan and Jeremy 
Zuppinger; Jo Daley; but above all, the brave villagers of San Cristobal and Daniel La 
Torre Lopez, the bear-hunter turned guide. 


References: 

Collar, N. J., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to watch 2—the world list of 
threatened birds. BirdLife International, Cambridge, U.K. 

Davies, C. W. N., Barnes, R., Butchart, S. H. M., Fernandez, M. & Seddon, N. 1994. 
The conservation status of the Cordillera de Colan. Unpublished report available from 
C. Davies at the address below. 

Fjeldsa, J. 1994. Geographical patterns for relict and young birds in Africa and south 
America and implications for conservation priorities. Biodiversity and Conservation 
Vol. 3 No. 3: 207-266. 

Fjeldsa, J. & Krabbe, N. 1990. Birds of the High Andes. (Copenhagen and) Svendborg: 
Zoological Museum, University of Copenhagen and Apollo Books. 

Hilty, S. L. & Brown, W. L. 1986. A Guide to the Birds of Colombia. Princeton Uniy. 
Press. ) 
Ridgely, R. S. & Tudor, G. 1994. A Guide to the Birds of South America. Vol. 2, the 

sub-oscine passerines. Oxford Univ. Press. 

Seddon, N., Barnes, R., Butchart, S. H. M., Davies, C. W. N. & Fernandez, M. 1996. 
Recent observations and notes on the ecology of the Royal Sunangel Heliangelus 
regalis. Bull. Brit. Orn. Cl. 116: 46-49. 

Snow, D. 1982. The Cotingas: bellbirds, umbrellabirds and their allies. British Museum 
(Natural History) and Oxford Univ. Press. 

Stattersfield, A. J., Crosby, M. J., Long, A. J. & Wege, D. C. 1995. A Global Directory 
of Endemic Bird Areas. BirdLife International, Cambridge, U.K. 


Addresses: Roger Barnes, 4 Claremont Drive, Leeds LS64ED, U.K. Stuart H. M. 
Butchart, Dept. of Zoology, Downing Street, Cambridge CB2 3EJ, U.K. Charles 
W.N. Davies, c/o 53 New Dover Road, Canterbury, Kent CT1 3DD, U.K. Mirko 
Fernandez, c/o APECO, Parque José de Acosta 187, Magdelena del Mar, Lima, Peru. 
Nathalie Seddon, Dept. of Zoology, Downing Street, Cambridge CB2 3EJ, U.K. 


© British Ornithologists’ Club 1997 


In Brief 75 Bull. B.O.C. 1997 117(1) 


IN BRIEF 
A SUBSTITUTE NAME FOR THE BIOKO RACE OF PYCNONOTUS VIRENS 


In a recent letter Mr R. J. Dowsett reminded me that Andropadus, 
which was long combined in the genus Pycnonotus, has again been 
recognized as a distinct genus (cf. Keith, Urban & Fry 1992, Birds of 
Africa Vol. 4: 287). Thus Pycnonotus virens poensis Dickerman, Bull. 
Brit. Orn. Cl. 114: 276, was preoccupied even when published by 
Stelgidillas poensis Alexander (Sipopo, Fernando Po). Stelgidillas 
Oberholser 1905 (Proc. U.S. Nat. Mus. 22: 30), the type of which is 
Andropadus gracilirostris, is a synonym of the latter genus. Therefore I 
propose the substitute name: Andropadus virens amadoni, nom. 
nov. Replaces Pycnonotus virens poensis Dickerman, 1994, not 
Stelgidillas poensis Alexander, 1903, Bull. Brit. Orn. Cl. 13: 35 
(now=Andropadus gracilirostris poensis). 

The name recognizes the contribution of Dr Dean Amadon to our 
knowledge of the birds of the Gulf of Guinea. 


American Museum of Natural History, ROBERT W. DICKERMAN 
Central Park West at 79th Street, 

New York, NY 10024, 

U.S.A. 29 April 1996 


THE THREAT STATUS OF THE SIDAMO LARK 


Good as it was to read Iain Robertson’s account of his observations of 
the Sidamo Lark WHeteromirafra sidamoensis in November 1994 
(evidently not 1974, as first indicated in his article) (Bull. Brit. Orn. Cl. 
115: 241-243), he is mistaken in regarding the ‘“‘Indeterminate”’ status 
assigned to this species in Threatened Birds of Africa (1985) as 
inconsistent. He bases this view on the fact that two other endemics of 
southern Ethiopia were classified as “‘Rare’’ despite their larger ranges 
and greater proportion of available habitat. I suspect the confusion 
arises because the word “‘indeterminate’’ suggests less urgency than the 
word “‘rare’’. 

The categories used in Threatened Birds of Africa, which were those 
then formally adopted by IUCN for all threatened species classifi- 
cation, are explained in the Introduction (pp. xxv—xxvi). I quote: 
““Indeterminate’ applies to ‘taxa known to be Endangered, Vulnerable 
or Rare but where there is not enough information to say which of the 
three categories is appropriate’ (... it should be noted that, since an 
Indeterminate species can be at best Rare, Indeterminate is a category 
of threat higher than Rare).”’ 

The assignment of Indeterminate to the Sidamo Lark in 1985 was 
made in the knowledge of John Ash’s 1974 observation, since he had 
kindly furnished an early draft of the paper he published in Bull. Brit. 
Orn. Cl. (105: 141-143) in 1985, but also in the light of his in litt. 


In Brief 76 Bull. B.O.C. 1997 AGT (#) 


comment, which I reproduced, that there was then “plenty of 
apparently suitable habitat’? in the area. Following his discovery in 
1989 that both sites at which the species had previously been recorded 
were being affected by man (Scopus 13: 90-97), and with the 
reformulation of the IUCN categories, the Sidamo Lark was classified 
as Endangered two years ago (Collar et al. 1994, Birds to Watch 2, 
which see also for the new categories). 

It is encouraging to know that some of these human influences 
appear now to have ceased, but I cannot think that such news will make 
a difference to the 1994 classification of the species. This is, however, a 
notable instance of the difficulty of appropriate categorisation in the 
absence of data. If it is accepted that highly cryptic birds in relatively 
poorly known regions stand a reasonable chance of being found over 
much wider ranges than those reported, then the Sidamo Lark might 
better have been classified “‘Data Deficient’’, which stands outside the 
new threatened categories. However, given the propensity for birds in 
Ethiopia and Somalia to be confined to often inexplicably small ranges, 
the precautionary principle requires us to entertain the possibility that 
the entire range of the species may well not extend beyond the small 
area south of Negele in which it has so far been found. It therefore 
appears that Iain Robertson and I have been and remain in closer 
agreement over the threat status of the Sidamo Lark than he previously 
imagined. 


BirdLife International, N. J. COLLAR 
Wellbrook Court, 

Girton Road, 

Cambridge CB3 ONA, 

UL 8 February 1996 


RANGE EXPANSION AND SUMMERING OF PALM WARBLER DENDROICA 
PALMARUM IN VENEZUELA 


On 29 June 1995, the first author observed a Palm Warbler Dendroica 
palmarum during an ornithological survey around the Mucubaji Lake, 
Mérida State in Venezuela (08°48'N, 70°48’'W), at an elevation of 
3500 m. The bird was also seen by several bird watchers and a 
photographic record was taken. Only one individual was seen for a few 
minutes, feeding on the ground and continually wagging its tail. The 
area is Open with very few trees, the dominant plant being Espeletia 
(Espeletia spp.). 

The solid rufous cap and the combination of yellow undertail 
coverts, pale yellow throat and whitish, vaguely streaked underparts 
differentiates this species from any other migrant parulid. No resident 
species are similar in colour. This coloration is typical of breeding adult 
plumage and distinctive of the nominate subspecies D. p. palmarum. 
The Palm Warbler breeds in North America and winters on the Gulf 
and southern Atlantic U.S. coasts, in eastern Yucatan, and in the West 


ia Tenefe 77 Bull. B.O.C. 1997 117(1) 


Indies; it has been occasionally recorded in Antioquia, Colombia, in 
October, and three times in Curacao and Aruba in February, March 
and November (Curson et al. 1994, Voous 1983). The nominate 
subspecies breeds in the western part of the North American range. 

It is well known that the Mucubaji area and the Chama River 
Canyon in Mérida State are important routes for several species of 
northern migrants (Phelps 1961). Our June observation suggested that 
this individual stayed on the wintering grounds during the following 
summer. This is apparently rare in warblers. There is one record of 
Blackpoll Warbler Dendroica striata from Chile in June 1858 (Philippi 
1940), and of Black and White Warbler Mniotilta varia and Northern 
Waterthrush Sezurus noveboracensis from the Netherlands Antilles in 
June (Voous 1983). 


We are grateful to Brian G. Prescot for providing the photographic record of our 
observation, and to Raymond McNeil and Clemencia Rodner for their suggestions and 
comments on this manuscript. 


References: 

Curson, J., Quinn, D. & Beadle, D. 1994. Warblers of the Americas. An Identification 
Guide. Houghton Mifflin Co., Boston. 

Phelps, W. H. 1961. Night migration at 4,200 meters in Venezuela. Auk 78: 93-94. 

Philippi, R. A. 1940. Aves migratorias Norteamericanas que visitan Chile. Bol. Museo 
Nac. Hist. Nat. Santiago 18: 65-85.. 

Voous, K. H. 1983. Birds of the Netherlands Antilles. 2nd edn. Van Dorp-Eddine n.v., 
Curacao. 


Sociedad Conservacionista Audubon 


de Venezuela, GUSTAVO RODRIGUEZ 
Aptdo. 80450, Caracas 1080-A. 

Coleccion Ornitologica Phelps, MIGUEL LENTINO 
Aptdo. 2009, Caracas 1010-A, 

Venezuela. 21 March 1996 


HISTORICAL RECORDS OF THREATENED AND NEAR THREATENED 
ARGENTINIAN BIRDS FROM MUSEO PROVINCIAL DE CIENCIAS 
NATURALES “FLORENTINO AMEGHINO” OF SANTA FE, ARGENTINA 


The Museo Provincial de Ciencias Naturales ‘‘Florentino Ameghino”’ 
in Santa Fe, Argentina, has a small but regionally important collection 
of 2300 bird specimens. It includes specimens of a number of species 
considered by Collar et al. (1992) to be threatened or near threatened, 
details of which are given below. Scientific nomenclature follows 
Altman & Swift (1993), English names Meyer de Schauensee (1982). 


Threatened species 


CROWNED EAGLE Harpyhaliaetus coronatus 


Two specimens: MFA-ZV-113 male, Campo La Eugenia (Dpt. San 
Cristobal, Prov. Santa Fe), 14 Jul 1957, R. Murnie coll. MFA-ZV-988 


In Brief 78 Bull. B.O.C. 1997 117(1) 


(sex?), Tintina (Dpt. Moreno, Prov. Santiago del Estero), without date, 
exchange with Escuela No. 62, collected before May 1950. 


BLACK-FRONTED PIPING-GUAN Pipile jacutinga 


MFA-ZV-1640 male, Km. 55, El Soberbio (Dpt. Guarani, Prov. 
Misiones), 24 Sep 1971, C. Rios coll. 


SPECKLED CRAKE Coturnicops notata 


MFA-NV-1825 female, Piquete (Dpt. La Capital, Prov. Santa Fe), 
21 Jan 1941, R. Muhn coll. 


HELMETED WOODPECKER Dryocopus galeatus 


MFA-ZV-1606 female, Dos de Mayo (Dpt. Cainguas, Prov. 
Misiones) 18 Jun 1968, E. Maletti coll. 


STRANGE-TAILED TYRANT Alectrurus risora 

Three specimens: MFA-ZV-1113 male, MFA-ZV-1118 male, 
MFA-ZV-1119 female, all from Ita-Ibaté (Dpt. Gral. Paz, Prov. 
Corrientes), 28 May 1952, A. Bonetto-G. Martinez Achenbach coll. 


MARSH SEEDEATER Sporophila palustris 


MFA-ZV-1985 male, Concordia (Dpt. Concordia, Prov. Entre Rios), 
13 Aug 1976, C. Garcia coll. 


Near threatened 


GREATER RHEA Rhea americana 

Three specimens: MFA-ZV-1431 young male, La Brava (Dpt. San 
Javier, Prov. Santa Fe), 18 Dec 1961, G. Martinez Achenbach coll.; 
MFA-ZV-1878 female, V. Saralegui (Dpt. San Cristobal, Prov. Santa 
Fe), 30 Nov 1948, H. Gavarro coll.; MFA-ZV-1161 young male, Santa 
Fe (Dpt. La Capital, Prov. Santa Fe), 26 Dec 1952, I. Costa coll. 


SOLITARY TINAMOU Tinamus solitarius 


MFA-ZV-1987 female, Cerro Moreno (Dpt. Cainguas, Prov. 
Misiones), 2 Aug 1978, C. Rios coll. 


HARPY EAGLE Harpia harpyja 


(1), MFA-ZV-2020 (sex?), Cerro Moreno (Dpt. Cainguas, Prov. 
Misiones), without date, was collected by E. Maletti before Oct 1980. 


HUDSONIAN GODWIT Limosa haemastica 


MFA-ZV-1460 female, Recreo, Salado River swamps (Dpt. La 
Capital, Prov. Santa Fe), 1 Feb 1948, G. Martinez Achenbach coll. 


RED-CAPPED PARROT Pionopsitta pileata 


Two specimens: MFA-ZV-268 male, MFA-ZV-267 female, from El 
Cerrito Island (Dpt. Bermejo, Prov. Chaco), 16 Jul 1944, C. Rios coll. 


In Brief 79 Bull. B.O.C. 1997 117(1) 


BLACK-BODIED WOODPECKER Dryocopus schulzi 


Two specimens: MFA-ZV-1614 male, Canada Ombt (Dpt. Gral. 
Obligado, Prov. Santa Fe), 15 Jul 1970, C. Rios coll.; MFA-ZV-1497 
male, Colonia Castelli (Dpt. Gral Guemes, Prov. Chaco), 27 Aug 1963, 
C. Rios coll. 


SHEAR-TAILED GRAY TYRANT Muscipipra vetula 


MFA-ZV-1996 male, Salto Encantado (Dpt. Aristdbulo del Valle, 
Prov. Misiones), 29 Jul 1978 C. Rios coll. 


BLACK-MASKED FINCH Coryphaspiza melanotis 


Four specimens: MFA-ZV-1670 male, Carabajal Island (Dpt. La 
Capital, Prov. Santa Fe), 10 Sep 1944; MFA-ZV-1359 male, Riacho 
fata re (Dpt.  La’*Capial; “Prov Santa” Fe), © 30° "Ade “1959; 
MFA-ZV-639 (sex?), Carabajal Island, 10 Jun 1945; MFA-ZV-638 
(sex?), 20 May 1944, Arteaga Island (Dpt. La Capital, Prov. Santa Fe), 
20 May 1944; all collected by G. Martinez Achenbach. 


CHESTNUT SEEDEATER Sporophila ruficollis 


Three specimens: MFA-ZV-619 (two males and one female with the 
same number), Santa Fe (Dpt. La Capital, Prov. Santa Fe), 10 Oct 
1933, M. Tesera coll. 


We thank Prof. Edelvita Fioramonti and Lic. Carlos Virasoro of Museo Provincial de 
Ciencias Naturales “Florentino Ameghino’”’ and M. Sc. Adolfo Beltzer of Instituto 
Nacional de Limnologia (CONICET), Santa Fe. 


References: 

Altman, A. & Swift, B. 1993. Checklist of the Birds of South America. 3rd edn. 
Book Masters, Inc., Ashland, Ohio. 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
T. A. & Wege, D. C. 1992. Threatened Birds of the Americas. The ICBP/IUCN Red 
Data Book. 3rd edn, pt. 2. International Council for Bird Preservation, Cambridge. 

Meyer de Schauensee, R. 1982. A Guide to the Birds of South America. Intercollegiate 
Press Inc. 


Instituto Nacional de Limnologia (INALI-CONICET), M. ORDANO 
Jose Macia 1933, 

(3016) Santo Tomé, Santa Fe, 

Argentina. 

Museo Provincial de Ciencias Naturales A. BOSISIO 
“Florentino Ameghino’’, 

Primera Junta 2859, 

(3000) Santa Fe, 

Argentina. 10 April 1996 


Books Received 80 Bull. B.O.C. 1997 117(1) 
BOOKS RECEIVED 


Lambert, F. & Woodcock, M. 1996. Pittas, Broadbills and Asities. Pp. 271, 24 colour 
plates, maps, text-figures. Pica Press. ISBN 1-873403-24-0. £26. 24 x 16 cm. 

Restall, R. 1996. Munias and Mannikins. Pp. 264, 80 colour plates, maps, text-figures. 
Pica Press. ISBN 1-873403-51-8. £28. 24 x 16 cm. 


The two latest in the Pica Press guides to bird families (or parts of families). The 
essential character of this outstanding series will doubtless be known to readers. Here, in 
the limited space available, it seems best to mention only the more particular features of 
the two under review. 

Frank Lambert’s account of two little known families, which are not only fascinating in 
themselves but important in our understanding of avian biogeography, aims to be fully 
comprehensive, and makes use of much unpublished data as well as his own field research 
over many years, together with a thorough study of museum specimens and the previous 
literature. The numerous unpublished observations by others are all acknowledged with 
the observer’s name. It thus provides the essential base on which all future studies will 
stand, the last monographic treatment of either family being Elliot’s Monograph of the 
Pittidae (2nd edn, 1893). The extraordinary courtship behaviour, still hardly understood, 
of a bird such as the Green Broadbill should surely spur young ornithologists to take 
up the challenge of documenting and filming the life history of this and many of the 
other species described here. The colour plates by Martin Woodcock, illustrating all the 
species and the main subspecies, in their various plumages, are spectacular and 
beautifully reproduced. 

Robin Restall’s book is a one-man tour de force. It deals with one section of the 
Estrildidae which (although split by some authors) he considers best treated as a single 
enlarged genus, Lonchura, with 40 known species. The text, which follows the standard 
pattern for books in the series, is packed with unpublished information and is illustrated 
with a large number of the author’s excellent, lively drawings of display postures, as well 
as diagrammatic figures of mouth markings of nestlings. Of the colour plates, the first 16, 
placed before the main text, illustrate all the species and well-marked subspecies in 
standard poses, Plates 17-80, which come at the end of the book as a sort of appendix, are 
a superb collection, of Tunnicliffe quality, of the author’s paintings from the life of 
individual birds of most of the species, most of them shown with a wing extended, from 
above and below, as well as in side view; all are accompanied by hand-written notes on 
points of colour, feather pattern, sex or geographical differences etc. ‘Those interested in 
scientific bird painting as an art will want to buy this book for these 64 plates alone. 


Roselaar, C. S. 1995. Songbirds of Turkey: an atlas of biodiversity of Turkish passerine 
birds. Pp. 240, 145 maps. Pica Press. ISBN 90-74345-07-7. £24. 24 x 17 cm. 


This work, the result of more than 20 years’ research (much of it summarised in BWP), 
is essential for anyone interested in the subspecific variation of west Palearctic birds. 
Turkey is in a key position in the west Palearctic, with its connections to southeast 
Europe, the Caucasus area to the east and the Near East to the southeast, and its 
Mediterranean coastal habitats in the south. It was not possible to include in BWP all the 
details relevant to a full account of the situation in Turkey; this is presented here, 
together with the most complete distribution maps available to date. Records are mapped 
individually, not by grid squares, an essential requirement for the detailed analysis that is 
presented and discussed. 


Sea Swallow (Annual Report of the Royal Naval Birdwatching Society), vol. 45, 1996. 
Pp: 120) 


Volume 45 of Sea Swallow is an enlarged Golden Jubilee edition, celebrating the first 
50 years of the Society’s life since its foundation just after the end of the last war. In 
addition to reports and papers of the usual kind, it includes a number of interesting 
reviews of the development of birdwatching at sea, from the earliest days to the present. 
There is also a special 8-page colour section of RNBWS photographs depicting 26 
species, some of them sea-birds and some ship-borne migrants. Non-members can obtain 
a copy (price £8, including postage in U.K.) from the Editor, our Hon. Secretary. 


NOTICE TO CONTRIBUTORS . 
Papers, from Club Members or non-members, should be sent to the Editor, Dr 
D. W. Snow, The Old Forge, Wingrave, Aylesbury, Bucks HP22 4PD, U.K., 
and must be offered solely to the Bulletin. They should be typed on one side of 
the paper, with treble-spacing and a wide margin, and submitted in duplicate. 
The style and lay-out should conform with usage in this or recent issues of the 
Bulletin. 


A contributor is entitled to 10 free offprints (16 if 2 or more authors) of the pages of the 
Bulletin in which his contribution, if one page or more in length, appears. Additional 
offprints or offprints of contributions of less than one page may be ordered when the 
manuscript is submitted and will be charged for. Authors may be charged for proof 
corrections for which they are responsible. 


MEMBERSHIP 
Only Members of the British Ornithologists’ Union are eligible to join the 
Club, and to receive (postage free) four quarterly issues of the Bulletin, and the 
annual index, for an annual subscription of £12 (or U.S. $26). Applications, 
enclosing the annual subscription, should be made to the Hon. Secretary 
(address as below). 


NON-MEMBER SUBSCRIBERS & APPLICATIONS FOR BACK 
NUMBERS OR OTHER PUBLICATIONS 

The Bulletin (for 1997 onwards), together with annual index, may be 
purchased (postage free) by non-members on payment of an annual sub- 
scription of £22 (or U.S. $45) on application to The Publications Officer, S. J. 
Farnsworth, Hammerkop, Frogmill, Hurley, Maidenhead, Berks SL6 5NL, 
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BOC Occasional Publications series may similarly be obtained, on request 
to him. 3 


PAYMENTS 

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Payments in other currencies must include a further £4 for UK bank charges 
(except for annual rates in U.S. dollars, which are inclusive). All cheques or 
drafts should be made payable to the British Ornithologists’ Club. If preferred, 
remittances may be made by bank transfer direct to the Club’s bank 
account—Barclays Prime Account, Dale House, Wavertree Boulevard, 
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Road, Bickley, Bromley, Kent BR1 2LF. 


CORRESPONDENCE 
Correspondence on membership, changes of address and all other matters 
should be addressed to the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA, U.K. For details 


of Club Meetings see inside front cover. 


' Registered Charity No. 279583 


CONTENTS 


CLUB NOTICES Report of the Committee, Annual General Meeting, Meetings . 
GOODMAN, S. M., HAWKINS, A. F. A. & DOMERGUE, C. A. A new species of vanga 
(Vangidae, Calicalicus) from southwestern Madagascar............... 
BORGHESIO, L. Observations on the ecology of Tauraco ruspolii and T. leucotis in 
southern Ethiopia . : os... sae a ee aes & ene ce ee 
GOMEZ DE SILVA G., H. Further observations on the nesting of the Azure-rumped 
ERICSON, P. G. P. Swedish records of the eastern Palearctic Hoopoe subspecies 
Upupa epops saturata...\.o0.. 3 on ee Se oe eee 
GALETTI, M., MARTUSCELLI, P., PIZO, M. A. & SIMAO, I. Records of Harpy and 
Crested Eagles in the Brazilian Atlantic forest... .. . . 9. a ee 
BROOKS, T. & DUTSON, G. ‘I‘wenty-nine new island records of birds from the 
Philippines... 2. ge ee PRP a ee Oe 
FRITH, C. B. & FRITH, D. W. ‘The taxonomic status of the bird of paradise 
Paradigalla carunculata intermedia (Paradisaeidae) with notes on the other 
Paradigalla taxa io. \36 6 ce 8 Paes io Ss ee 
DOWSETT, R. J. & MOORE, A. Swamp warblers Acrocephalus gracilirostris and 
A. rufescens at Lake Chad;-Nigeria... 0): 1... . :).).4 2 ee ee 
CLANCEY, P. A. Variation in the Cape Penduline Tit Anthoscopus minutus of the 
southern Afrotropics 2!) 3)../2°. 1.9 sie Di. Sa oe eee ee 
HAZEVOET, C. J. Ona record of the Wattled Crane Bugeranus carunculatus from 
Guinea-Bissau. 2 cay WR a we a ee oe 
ERICSON, P. G. P. & AMARILLA, L. A. First observations and new distributional data 
for birds in Paraguay ie. o226/s. 0. ees Yo We 
MEES, G. F. On the identity of Heterornis senex Bonaparte ................ 
BARNES, R., BUTCHART, S. H. M., DAVIES, C. W. N., FERNANDEZ, M. & SEDDON, N. 
New distibutional information on eight bird species from northern Peru . . 
In Brief DICKERMAN, R. w. A substitute name for the Bioko race of Pycnonotus 
VITENS. ics ne SR Ce hw Ripe es a 5 ee 
COLLAR, N. J. The threat status of the Sidamo Lark...) 23 3g 
RODRIGUEZ, G. & LENTINO, M. Range expansion and summering of Palm 
Warbler Dendroica palmarum in Venezuela................. 
ORDANO, M. & BOSISIO, A. Historical records of threatened and near 
threatened Argentinian birds from Museo Provincial de Ciencias 
Naturales “‘Florentino Ameghino” of Santa Fe, Argentina...... 
BOOKS RECEIVED. . «00006060564 4,4 @ « «soe 4 eum © diese le mele ce eh 


76 


fis 
80 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 


mail, after their current subscription has been paid. 


COMMITTEE 
D. Griffin (Chairman) (1993) Miss H. Baker (1994) 
Revd T. W. Gladwin (Vice-Chairman) (1993) S. J. Farnsworth (1990) 
D. J. Montier (A/Hon. Treasurer) (1996) R. E. F. Peal (1993) 
Cdr M. B. Casement, OBE, RN Dr R. P. Prys-Jones (1995) 
(Hon. Secretary) (1996) N. H. F. Stone (1995) 


Hon. Editor: Dr D. W. Snow 
Chairman of Publications Sub-committee: Mrs A. M. Moore 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


ar 
Bs 
BeGs 


ISSN 0007-1595 
= D> 


Bulletin of the 


British Ornithologists’ Club 


Edited by 
Dr R. A. CHEKE and Dr D. W. SNOW 


Swit HSONIA/) 


JUL 2.9 199/ 


LIBRARIES 


Jolume 117 No. 2 June 1997 


FORTHCOMING MEETINGS 


Tuesday 16 September 1997. Graeme Green will speak on ‘‘Cotingas, and 
their niche in the neotropical avifauna’”’. Graeme has long had a special interest 
in the taxonomy, identification, ecology and conservation of tropical birds. He is an 
active member of the Oriental Bird Club and the Neotropical Bird Club, travelling 
widely in pursuit of these interests. He is co-author of Cotingas and Manakins, due 
to be published by Pica Press in 1998. 

Applications to the Hon. Secretary by 12 September, please. 


Tuesday 4 November 1997. Tony Marr will speak on “An Atlantic Seabird 
Odyssey’’. Tony has had a life-long interest in seabirds, especially through 
‘pelagics’ in the eastern Atlantic, and he has pursued them from a wide variety of 
boats and craft, off Portugal, ‘The Canaries, Madeira, West Africa, and the South 
Atlantic. Having retired early to devote more time to birds, he is well-known to 
many as a tour leader to many parts of the world, and as a lecturer and author of 
articles on seabird topics. He serves on the BOU Records Committee, and the 
Seabirds Advisory Panel for British Birds. 

Applications to the Hon. Secretary by 21 October (Trafalgar Day!), please. 


Meetings are held in the Sherfield Building of Imperial College, South 
Kensington, London SW7, at 6.15 p.m. for 7 p.m. The nearest Tube station is at 
South Kensington, and car parking facilities are available; a map of the area will be 
sent to members, on request. The cash bar is open from 6.15, and a buffet supper, of 
two courses followed by coffee, is served from about 7.00. (A vegetarian menu can be 
arranged if ordered at the time of booking.) Informal talks are given on completion. 


Overseas Members visiting Britain are particularly welcome at meetings. 
For details in advance, please contact the Hon. Secretary, Cdr M. B. 
Casement, OBE, RN, Dene Cottage, West Harting, Petersfield, Hants. 
GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1997 


Apart from single copies made for the purposes of research or private study, or 
criticism or review, as permitted under UK law, no part of this publication may be 
reproduced, stored or transmitted in any form or by any means, except with prior 
permission in writing of the publishers, or in accordance with the terms of licences 
issued by the Copyright Licensing Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; 
for address see inside back cover. 


81 Bull. B.O.C. 1997 117(2) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 117 No. 2 Published 20 June 1997 


The eight hundred and sixty-fifth meeting of the Club was held on Tuesday, 21 January 
1997, at 6.15pm. 32 Members and 15 guests attended. 

Members present were: D. GRIFFIN (Chairman), Professor C. J. FEARE (Speaker), Miss 
H. Baker, J. W. BaRRINGTON, Captain M. K. Barritt RN, P. J. BELMaN, I. R. BisHop, 
Mrs D. M. Braptey, D. R. CaLper, Cdr M. B. CasEMENT RN, Professor R. J. 
CHANDLER, Dr R. A. CHEKE, S. J. FARNSWORTH, Rev. T. W. GLADWIN, Dr A. GOSLER, 
C. A. R. Heim, R. M. Kettiez, Dr C. F. Mann, D. J. MontigErR, R. G. Morcan, Mrs 
A. M. Moors, Mrs M. N. MuLter, Dr R. P. Prys-Jonss, N. J. REDMaN, S. J. R. RUMsEy, 
R. E. Scott, P. J. SeLuar, S. A. H. StatHam, Dr D. W. Snow, N. H. F. SToneE, Mrs 
F. E. Warr, Professor W. E WarTERS. 

Guests attending were: Dr E. GiLu (Speaker), Mrs S. GrirFin, Miss G. BONHAM, 
M. Braptey, Mrs J. B. CaLper, Mrs C. R. CASEMENT, Mrs J. M. GLapwin, A. MartTIN, 
Mrs M. Montier, P. J. Moore, M. Paine, R. Ranft, Mrs B. K. Snow, Cdr J. M. W. 
Tope RN, J. Warr. 

After dinner, Professor Chris Feare and Dr Elaine Gill gave a joint presentation under 
the title ““The Biology of Pink Omelettes’’, illustrated by a splendid series of their own 
slides. The following is a brief synopsis: 

During June and July, omelettes and other culinary treatments of the pink-yolked eggs 
of Sooty Terns Sterna fuscata provide the people of the Seychelles with a traditional and 
much sought-after delicacy. The eggs are harvested from large colonies, mainly on 
remote islands, and transported by boat to the main island, Mahé, for sale and 
distribution. Since the 1940s, concern has been expressed about over-exploitation, 
following declines in colony size and some colony extinctions. Although many of these 
problems have in fact been caused by habitat changes on breeding islands, the Seychelles 
Government wants to ensure that exploitation is sustainable and is therefore supporting 
research on the biology of the Sooty Tern and the management of the islands where they 
breed. 

Sooty Terns normally lay their single egg in a scrape on open, lightly-vegetated 
expanses on flat islands; in Seychelles they also nest on rocky islands and, exceptionally, 
on the floor of woodland. The breeding season lasts from June to October, each pair 
taking around three months to raise its offspring to fledging. During studies in the early 
1970s, under an NERC grant to Prof. George Dunnet, it was discovered that egg-laying 
within a colony is highly synchronous, with most eggs being laid within nine days. The 
fledging success of eggs laid after the period of intense laying early in the season declines 
markedly. After losing an egg, parents may return to lay a replacement about 13 days 
later, but the ability to replace lost eggs also shows a marked seasonal decline. 

The current studies by both speakers were initially funded by the Royal Society, 
Seabird Group, Percy Sladen Memorial Fund, Bird Island Lodge and Air Seychelles, 
but are now largely supported by the Seychelles Government, Island Development 
Corporation and Bird Island Lodge. The aims are to investigate inter-colony movements, 
through ringing and inter-genetic studies; to determine from ringed birds their age at first 
breeding and longevity; and to assess the potential effects of island vegetation 
management on nest density and colony size. Thereafter the aim is to establish schemes 
for monitoring colony size, breeding success and egg harvests. Using these and other 
data, the intention is to produce a model of Sooty Tern population dynamics, from which 
a sustainable egg harvest and harvest strategy can be derived. 

A regulated harvest of Sooty Tern eggs ensures that the exploited colonies are 
protected during the laying season. This prevents poaching, which can be destructive on 
unprotected islands, and also confers other benefits on Sooty Tern islands—for example 
the protection from illegal killing of other species, such as Masked and Brown Boobies 
Sula dactylatra and SS. leucogaster, and Green Turtles Chelonia mydas. All of these have 
suffered serious declines in the Seychelles, despite being legally protected, but while 
oy Tern eggs are being collected the legal protection afforded to these species can be 
enforced. 


Meetings 82 Bull. B.O.C21997 Mia?) 


The eight hundred and sixty-sixth meeting of the Club was held on Tuesday, 18 
March 1997, at 6.15pm. 21 Members and 8 guests attended. 

Members present were: D. GRIFFIN (Chairman), Miss H. Baker, J. W. BARRINGTON, 
P. J. BELMan, Cdr M. B. CasEMENT RN, Professor R. J. CHANDLER, A. Gripps, Rev. 
T. W. Grapwin,. J. A. Jopitine, C. A. R. HELo, Lt. Col. P. S. Kerr-Smitey, Dr J. F. 
Monk, D. J. Montrer, R. G. Morcan, Mrs A. M. Moore, Mrs M. N. MULukr, R. C. 
Price, Dr W. G. Portsous, N. H. F. STONE, Mrs F. E. Warr, Professor W. E. WaTERs. 

Guests attending were: Major F. CouRTENAY-THOMPSON (Speaker), Mrs C. R. 
CASEMENT, Miss F. CoURTENAY-THOMPSON, Dr D. Foskett, Mrs J. M. GLapwin, Mrs 
M. Montisr, P. J. Moore, J. Warr. 

After dinner Major Frank Courtenay-Thompson spoke on the subject of ““Three Years 
Birdwatching in Saudi Arabia’’, illustrated with a wide range of colour slides. In 1970, 
when he arrived in Saudi Arabia, there were few good roads and travel was slow. Living 
in Jeddah, most of his bird watching was in the western region. The fascination was the 
geographical location and the variety of habitats, but the challenge was the lack of 
background information. 

Much has been published about the birds of Saudi Arabia in recent years, but 25 years 
ago there was very little. Meinertzhagen’s Birds of Arabia, published in 1954, was already 
out of print and scarce. The 1972 Collins Field-guide was the first to include the Middle 
East with Europe, and even that did not give the full picture. Frank published his own 
bird list in 1972, with the aim of helping people out there, rather than contributing to 
world knowledge. Copies were, however, circulated in Europe and America, and aroused 
some interest at the time. 

The west coast has many creeks and mangroves. A mountain range rising to over 
10,000 feet runs the length of the peninsula, on the southern half of which are extensive 
juniper and olive woodland. East of this range, a plateau extends to the Persian Gulf, and 
the two sand seas are centrally placed in the north and south. Seasonal temperature on the 
plateau varies from 120°F to below freezing, with a far smaller range and very high 
humidity at the coast. Much of the country has a brief period of rain most years, which 
can reach 20 inches in the mountains. 

Any green area becomes a haven for exhausted Palearctic-African migrants crossing 
Saudi Arabia but, following the blistering summer, there are less of these and a number 
of migrant species are only seen in the spring. Many are unfamiliar to the British 
ornithologist, the most colourful of which are the European Bee-eater Merops apzaster, 
Golden Oriole Oriolus oriolus and large parties of Hoopoes Upupa epops, some of which 
nest. There are twelve species of wheatear and six races of yellow wagtail. Flocks of 
European Cranes Grus grus fly over, often after dark. 

The southern tip of Arabia was once joined to Africa, and there is a remnant 
population of African (Ethiopian) origin. Typical of these are the Bateleur Eagle 
Terathopius ecaudatus, the Grey Hornbill Tockus nasutus, Ruppell’s Weaver Ploceus 
galbula and Abyssinian Roller Coracias abyssinica. Along the coast are to be found the 
Reef Heron Egretta gularis, Goliath Heron Ardea goliath and Little Green Heron 
Butorides striatus, as well as the Crab Plover Dromas ardeola. There is no evidence that 
any of these migrate to and from Africa, but they do move up and down their corner of 
the peninsula following the rains. 

There are eleven species which are only found in Arabia. These include the Arabian 
Babbler Turdoides squamiceps, White-eyed Gull Larus leucophthalmus and the 
increasingly rare Arabian Rock Partridge Alectoris philbyi and Arabian Woodpecker 
Dendrocopos dorae. In addition, a number of species are strictly birds of the desert belt, 
notably the Trumpeter Bullfinch Bucanetes githagineus, Isabelline Shrike Lanzius 
isabellinus and the Isabelline Wheateater Oenanthe isabellina. All appear as washed-out 
versions of brighter cousins. 

Saudi Arabia has a wide range of both migrant and resident species, but unlike most 
other parts of the world, is still largely unexplored by ornithologists. 


R.A. Cheke 83 Bull. B.O.C. 1997 117(2) 


AVIAN TAXONOMY FROM LINNAEUS 
| TO DNA 


PAPERS PRESENTED AT A JOINT MEETING 
BETWEEN THE BRITISH ORNITHOLOGISTS’ CLUB 
AND THE LINNEAN SOCIETY OF LONDON HELD 

AT BURLINGTON Housg, 23 Marcu 1996 


PREFACE 
by Robert A. Cheke 


The use of the binomial system to classify plants and animals, as 
formalised by Linnaeus, continues to be the means by which species are 
named. Acceptance of binomials after much controversy was followed 
by further, often heated, debates about the usefulness of the trinomial 
system and the idea of subspecies. The latter arguments occupied the 
members of the British Ornithologists’ Club (BOC) for many years, 
especially at the end of the 19th century. We have now come full circle 
and, once again, the subspecies concept is under criticism, but mostly 
for quite different reasons from those advanced by the contemporaries 
of the founders of the BOC. It is salutary that there is probably now 
more disagreement over what is meant by species than there has ever 
been (Malliet 1995). 

The purpose of the gathering, fittingly held in the meeting rooms 
of the Linnean Society of London, was to provide an opportunity to 
discuss modern views of avian taxonomy, at the same time 
remembering the historical context. The papers from the meeting 
which are published in full here are only those which addressed species 
concepts per se. Abstracts are included for three of the others, as well as 
a paper on Linnaeus’ correspondence with Scopoli, based on a poster 
presentation. 

Current arguments are not without practical import, and the 
consequences that their resolution might have, for purposes such as 
conservation or zoo-archaeology, was another major issue _ for 
discussion, as was the whole construct of traditional classification. If a 
population is genetically distinct, such as one of the groups of Red Kite 
Milvus milvus described by Parkin, why shouldn’t it be classed as a 
subspecies or even a species? Indeed, where can the line be drawn, 
since each individual bird has a unique genetic code? Does the logic 
necessitate the acceptance of many more species of birds than are 
hitherto recognised? Are traditional higher order groupings redundant? 

As Jeremy Greenwood points out in the introductory contribution, 
there are many taxa of taxonomic thought and each might classify the 
same group of animals differently. The abolition of subspecies is one 
consequence of the phylogenetic species concept (PSC) advocated by 
Robert Zink, whose conclusions would have been heard with approval 


R. A. Cheke 84 Bull. B.O.C. 1997 117(2) 


by some of the BOC’s founders, as well as by a few of the Italian 
ornithologists discussed by Violani and Barbagli. Zink’s arguments 
include conclusions derived from DNA data and analyses which would 
be difficult to obtain without the use of microprocessors. Although the 
post-Watson and Crick era has seen extraordinary advances in our 
understanding of biological processes at the molecular level, especially 
when coupled with information technology, DNA may yet only serve 
taxonomy as another set of “‘characters’’, so far as bird classifications 
are concerned. David Snow maintains support for the biological species 
concept and warns that the general adoption of the PSC could lead to 
decades of instability in taxonomy. He also draws attention to the 
proliferation of new species (though this in itself is not a valid argument 
against the PSC), which adherence to the adoption of the PSC would 
engender; and these two themes are taken up by Collar, who 
emphasises that the adoption of the PSC would make the role of 
international conservationists impossibly difficult. ‘The ‘problem’ of the 
recognition of extra species, under the PSC, was emphasised in an 
account of the meeting (Martin 1996) but disputed by Zink (1996), who 
pointed out that most of the putative extra species are already 
recognised as subspecies. 

Parkin demonstrated that there are genetic differences between 
populations of Red Kites from Germany, Spain and Wales, the latter 
being the least variable and having the poorest breeding success. 
Russell showed that it is possible to identify mummified birds of prey 
from X-rays, using taxonomic methods of much newer vintage than the 
mummies. The main conclusions reached in a study of eggshell 
structure to elucidate taxonomic relationships, principally at levels 
above the species (mainly sub-order or order), were summarised by 


Mikhailov. A detailed account of this research will appear elsewhere 
(Mikhailov 1997). 


References: 

Malliet, J. 1995. A species definition for the modern synthesis. TREE 10: 294-299. 

Martin, G. 1996. Birds in double trouble. Nature 380: 666-667. 

Mikhailov, K. E. 1997. Avian Eggshells: an Atlas of Scanning Electron Micrographs. 
British Ornithologists’ Club Occasional Publications series no. 3. Tring, U.K. (in 
press). 

Zink, R. M. 1996. Bird species diversity. Nature 381: 566. 


Address: Dr R. A. Cheke, Natural Resources Institute, University of Greenwich, 
Central Avenue, Chatham Maritime, Chatham, Kent ME4 4TB, U.K. 


© British Ornithologists’ Club 1997 


FJ. F. D. Greenwood 85 Bull. B:O.@>1997'117(2) 
Introduction: the diversity of taxonomies 


by Feremy 7. D. Greenwood 


Taxonomy and related fields are battle grounds onto which the 
non-combatant ventures at his peril, liable to be shot at from all sides. 
Even the definition of the subject is one on which its practitioners 
clearly disagree. I shall accept that: ““Taxonomy is, strictly speaking, 
the study of the principles and practice of classification’’ (Jeffrey 1977). 
Classifications have three main uses in biology: they allow us to 
summarise and organise our knowledge about living organisms, they 
help us to identify organisms, and they can provide an approximate 
summary of evolutionary relationships. The first is important because 
of the sheer diversity of living organisms and the second because that 
diversity makes identification difficult. Summarising evolutionary 
relationships is important because ‘‘Nothing in biology makes sense 
except in the light of evolution” (Dobzhansky 1973). So most biologists 
use classifications in most of their work. One might therefore assume 
that taxonomy would be a key element in the education of young 
biologists. In Britain, at least, this assumption would be completely 
unjustified; without having carried out any systematic investigation, I 
suspect that most undergraduate courses are devoid of formal teaching 
in taxonomy. As a result, most of us have a rather hazy knowledge of 
the principles and methods underlying classifications; even worse, 
because we use classifications every day of our lives, we may be 
unaware of quite how hazy our knowledge is. Furthermore, Arthur 
Cain’s (1959) prescient opinion that “‘we are about to see a considerable 
revision of the whole basis of taxonomic theory” has been amply 
justified; with the major developments in taxonomic philosophy, in 
sources of data, and in analytical methods that have occurred in the last 
four decades, the gap between the taxonomist and the users of the 
taxonomists’ products may, indeed, be wider than ever before. For that 
reason, I shall take some space to look at those developments, in the 
hope that my brief summary may be of use to others who feel the need 
to be more familiar with modern ideas in taxonomy but whose work, 
like mine, has prevented them from closely following those ideas as 
they have developed. My own recent education in the subject has relied 
particularly on: Ridley (1986), who provides a thought-provoking, if 
personal, view of the major schools of taxonomy from the point of view 
of an evolutionary biologist rather than of a practising taxonomist; 
Forey et al. (1992), who clearly explain cladistic views and methods; 
and Hillis & Moritz (1990), who cover many of the principles as well as 
covering in detail how molecular evidence may be used. (For those 
whose knowledge of molecular evolution is becoming rather rusty, Li & 
Graur (1991) provide a useful introduction to current ideas.) 


Organising knowledge 


We are able to use classification as a means of organising knowledge 
about living organisms particularly because species fall into clusters. 


J. J. D. Greenwood 86 Bull. B.O.C. 1997 117Q) 


a wi sshe C 


Figure 1. Clusters of species in a two-dimensional character space. Cluster A is distinct 
but do B and C represent one cluster or two? If they form two clusters, where should the 
dividing line be drawn? 


For example: bird species share certain characteristics in common, 
mammal species share other characteristics, and there is a gap between 
them; bats share some characteristics with birds but are still firmly 
linked to the mammals by most of their characteristics; bats do not 
cause us to have difficulty in recognising the two chief clusters of 
warm-blooded vertebrates or in distinguishing between them. We can 
make general statements about all members of such clusters, to help us 
reduce our knowledge base to manageable proportions. The process 
is further facilitated by the fact that we may arrange the clusters in 
a non-overlapping hierarchy, with clusters at each taxonomic level 
themselves being clustered at the level above. Unfortunately, the 
clusters into which species tend to fall are often indistinct (Fig. 1), 
especially when we consider fossils as well as extant species (Fig. 2), so 
the distinctions between taxa may not be clear and generalisations 
about the members of a taxon may not all apply to every species. 


Evolution: branching and divergence 


For many, the chief fascination of biology is that living organisms have 
an evolutionary history, being related to each other through descent 
from common ancestors. The history of life can be described by a 
simple branching pattern (e.g. Fig. 2) and that pattern can be reflected 
by the taxonomic hierarchy. Because evolution is central to biology, 
taxonomy has traditionally been used to summarise evolutionary 
relationships as well as to provide groups (clusters) about which general 
statements can be made. For example, as well as being seen to have 
many characteristics in common, species in the class Aves are 
recognised as sharing an evolutionary relationship closer than the 
relationship between any one of them and any species in other classes. 

If species that were evolutionarily closely related were always more 
similar than those that were more distantly related, classifications could 
easily reflect both degrees of similarity and degrees of relationship. 
Unfortunately, this would only be true if evolution involved species 
descended from a common ancestor simply becoming steadily more and 
more different from each other (Fig. 3a). Unfortunately, life is not that 
simple: rates of divergence vary, as in Fig. 3b, in which species 3 is now 


F. J. D. Greenwood 87 Bull. B.O.C.1997 117(2) 


1 
2 
F 3 
5 7 
et 5 g 
4 
5 
6 


time 
Figure 2. A hypothetical evolutionary tree in which six extant species (1-6) form two 


distinct clusters on the axis of divergence. The known fossils (7—9) link these clusters 
together. 


more different from species 2 than the latter is from species 1, even 
though species 2 and 3 share a more recent common ancestor; 
convergent evolution is equally destructive of the correspondence 
between similarity and evolutionary relationship (Fig. 3c). 


The diversity of taxonomies 


Because patterns of similarity and evolutionary relationships may not 
be congruent and because there are various ways of describing both 
similarities and relationships, different taxonomists may employ 
different principles and procedures in their work. For the purposes of 
exposition, I recognise five main groups: the traditional evolutionary 
taxonomists, pheneticists, distance-based evolutionary taxonomists, 
Hennigian cladists and pattern cladists. Cladists have dominated 
taxonomic thinking in recent decades and some (perhaps all) of them 
claim that the other schools are now extinct. That this is not true is 
shown by the fact that the best-known recent classification of birds 
(Sibley & Ahlquist 1990) is distance-based. Furthermore, many 
non-taxonomists have grown up knowing something about traditional 
evolutionary taxonomy and may assume that all biological classifi- 
cations rest on traditional principles and procedures. Indeed, because 
classifications take time to re-work, many of them still do. 

It is clearly important that those who use a particular classification 
should know and understand the taxonomic principles used to produce 
it. If they do not, they are likely to draw invalid conclusions from it. 
The most important message I have to deliver is that it is up to the 


F. J. D. Greenwood 88 Bull. B.O.C..1997 117(2) 


1 
CD) 
2 
S 
a 3 
2 gy) 
3 
1 
8 
(=| 
oO 
b eS 
3 2 
3 
1 
2 2 
S 
Cc i 
12 
ao} 
3 


SS SS 


time 


Figure 3. Three evolutionary trees. Note that each is meant to represent the actual course 
of evolutionary events, not what might necessarily be inferred from available evidence. In 
(a) species diverge at fairly steady rates, so that degrees of similarity between extant 
species reflect their evolutionary relationships. The correspondence between similarity 
and relationship is broken in (b) because rates of divergence differ and in (c) because of 
convergence. 


taxonomist to state clearly the principles and procedures involved in 
producing a classification and up to the user to pay proper attention to 
such statements. 


What sorts of characters to use? 


‘Taxonomists differ not only in their principles and procedures but also 
in the sorts of characters they use as the basis for their classifications. 
Morphological characters have traditionally been dominant but the use 
of other characters has a long history: not only have naturalists long 
used song to identify and distinguish similar birds (e.g. White 1789) 
but among formal taxonomists Linnaeus (1758) used behaviour 
(‘‘adscendit noctw’’, in respect of Lumbricus terrestris) and Nuttall (1904) 
used immunological characters. From time to time, individual taxo- 
nomists have promoted the use of particular sets of characters, either 
because of the ease with which they may be studied or because they are 


F. J. D. Greenwood 89 Bull. B.O.C. 1997 117(2) 


supposed to be particularly revealing of evolutionary relationships. The 
latter suppositions generally rely on questionable arguments about 
certain sorts of characters being evolutionarily conservative because 
they are likely to be less (or more!) subject to natural selection than 
other sorts. But, just as the history of evolutionary biology is littered 
with examples of characters once thought to be selectively neutral but 
now known to be strongly selected, so is the history of taxonomy 
littered with classes of characters no longer thought to be peculiarly 
valuable. Molecular (especially DNA) data are still often thought to be 
especially useful but they produce no more consistent pictures of 
phylogeny than do morphological data (Patterson et al. 1993, Sheldon 
& Bledsoe 1993). The best classifications are likely to result from 
considering all the available data—though for evolutionary and cladistic 
taxonomies homologous characters are of particular importance. 


Traditional evolutionary taxonomy 


What one regards as traditional evolutionary taxonomy is a matter of 
opinion. I refer to the approach, growing out of ‘““The Modern 
Synthesis’ of Stebbins (1950), Simpson (1961) and Mayr (1969), which 
has been further discussed by Cronquist (1988) and Bock (1989). It 
involves working out the evolutionary history of the species under 
consideration, taking into account evidence such as ecology and 
biogeography as well as the distribution of characters among species. 
Attention is paid to the function of characters, with assessments being 
made of the likelihood of different possible evolutionary changes; it is 
important to assess whether or not similar character states are homol- 
ogous or the result of convergence. This is because taxa are required to 
be monophyletic which means, for the traditional evolutionary 
taxonomist, merely that all group members should share a common 
ancestor, which should also be a member of the group (Fig. 4). Note 
that it is not necessary that all the descendants be included for a taxon 
to be regarded traditionally as monophyletic, so a divergent species (or 
group of species) can be separated from a group with which it shares 
common ancestry (as taxa VI and VII are separated from taxa III and 
IV in Fig. 4c). For the cladist (see below), in contrast, taxa III and IV 
(Fig. 4b and 4c) are paraphyletic (and not allowed); strict monophyly 
requires that all descendants are included in the group. Thus tra- 
ditional evolutionary taxonomies attempt to reflect both the branching 
pattern of the evolutionary trees and the extent of divergences. 

The problem with traditional evolutionary taxonomy is that it is 
highly subjective, both at the stage of working out the underlying 
evolutionary narrative and at the stage of converting the phylogenetic 
tree into a classification. 


Phenetic taxonomy 


Pheneticists dispense with the subjectivity of traditional evolutionary 
taxonomy by abandoning the attempt to summarise evolutionary 
relationships in the classification. For the phenetic school, taxonomy is 


JF. J. D. Greenwood 90 Bull. B.O.C. 1997 117(2) 


o 
=] 
a Sb 
5 
Es} II 
ii 
g 
S 
b 5 
oO 
> 
2 
IV 
I 
= VI 
5 
y bb vu 
oO 
> 
2 
IV 


time 


Figure 4. A hypothetical evolutionary tree and various classifications (a—c) for four extant 
species and two known fossils. Classification (a) would be acceptable to both traditional 
evolutionary taxonomists and cladists, since taxa I and II are monophyletic. Classification 
(b) would be more acceptable to a phenetic taxonomist, because taxon V reflects the 
similarity between the.two species falling into it; but this taxon is polyphyletic, so the 
classification would be rejected by both traditional evolutionary taxonomists and cladists. 
The latter would also object to the paraphyletic taxa III and IV and so would also object 
to classification (c), though this would be acceptable to traditional evolutionary 
taxonomists. Whether one of the latter preferred (a) or (c) would depend on the weight he 
or she gave to divergence relative to phylogenetic relationship. 


a matter of producing a hierarchy that reflects the inherent hierarchical 
clustering of nature. Species are grouped according to degrees of 
resemblance. In effect, species are seen as points in a multi-dimensional 
hyperspace, the dimensions corresponding to various characters and 
the positions along those dimensions being determined by how 
different the species are in respect of those characters. The phenetic 
taxonomist’s job is to establish the dimensions, to measure positions 
and distances, and to recognise clusters (and the clusters of clusters 
...). Thus the pheneticist would apply classification (b) in Fig. 4; the 
fact that taxon V is polyphyletic is immaterial, since the taxa are units 
of resemblance not of phylogeny. 

At first sight, phenetics is less subjective than traditional taxonomy 
because it requires no judgements about evolution. But, as Ridley 
(1986) and others have pointed out, it is possible to define and measure 


JF. F. D. Greenwood 91 Bull. BxOvC11997 T17(2) 


‘“‘similarity’ in a variety of ways and the choice between them is 
entirely arbitrary; furthermore, a large variety of different methods is 
available for defining clusters and the choice is again arbitrary. Because 
the classifications produced may differ markedly according to which 
similarity measures and clustering methods are used, it is difficult to 
argue that phenetic methods are superior to traditional methods. 


Distance-based evolutionary taxonomy 


The extent to which single-strand DNA from two different sources 
produces hybrid double strands under specified conditions depends on 
the extent to which the two DNAs contain identical sequences, so such 
hybridization provides a measure of similarity. If it were true that 
evolution at the level of DNA (base substitution) proceeded in a steady, 
undirected, clock-like manner and provided that one could properly 
correct for the likely occurrence of changes that limit divergence 
(parallel changes in the two species and multiple changes, including 
reversals, at single nucleotide sites), then the similarity between species 
would reflect their evolutionary relationship (as in Fig. 3a). An 
essentially phenetic classification based simply on the degree of 
resemblance (of DNAs) would then have an evolutionary interpret- 
ation. However, the speed at which the “molecular clock’’ runs is 
clearly far from constant (Hillis & Moritz 1990) and the corrections 
required for parallelism, multiple hits and reversals are based on 
somewhat arbitrary assumptions. Furthermore, distance measures 
based on DNA-DNA hybridization are subject to considerable 
experimental error (Werman et al. 1990). It is for these reasons and 
others that the classification of Sibley & Ahlquist (1990) aroused so 
much controversy (see Sheldon & Bledsoe 1993, Harshman 1994). It 
has, nonetheless, become widely accepted, as have other classifications 
based on the idea that the degree of similarity (especially of DNAs) can 
be used as a direct assessment of evolutionary relationships. 


Hennigian cladistics 


Hennig (1950, 1966) revolutionised systematics by proposing clear, 

non-arbitrary methods for exposing the patterns of diversity that result 

from phylogenetic branching and for describing those patterns in an 

hierarchical classification. Both Ridley (1986) and Forey et al. (1992) 

present clear introductions to cladistic methods. Forey et al. describe 

its axioms as: 

1. Nature’s hierarchy is discoverable and effectively represented by a 
branching diagram. 

2. Characters change their status at different hierarchical levels. 
Characters within a study group that are either present in all 
members of the study group or have a wider distribution than the 
study group (plesiomorphies) cannot indicate relationships within 
the study group. 

3. Character congruence is the decisive criterion for distinguishing 
homology (synapomorphy) from non-homology (homoplasy). 

4. The principle of parsimony maximises character congruence. 


J. F. D. Greenwood 92 Bull. B.O.C. 1997 117(2) 


character states 
cladogram I cladogram II 


2 Rt Bb CA ae ema es viiD: 
FB B 


A 3 rife BYE bie Ea 3 
FD CED 


oF BOT upeTie Ber ity OFF ST 


CE 


Figure 5. Two alternative cladograms for species 2, 3 and 4, based on six characters, for 
which the primitive states (seen in the ‘outgroup’ species 1) are represented by lower case 
letters and derived states by upper case letters. The dark bars represent points at which a 
primitive state changes to a derived state. Redrawn from Forey ez al. (1992). 


The branching diagrams produced by cladistic methods are referred to 
as Cladograms. By ‘“‘character congruence’”’ is meant the co-occurrence 
of characters, such that they specify the same taxonomic group. Thus 
in cladogram I of Fig. 5, C and E are congruent (since they both occur 
in, and only in, the group 3—4) whereas F is incongruent with them 
(since it occurs not only in some, but only some, members of the group 
but also outside the group). Incongruencies imply convergent 
evolution. Cladogram II has fewer congruencies and more incongru- 
encies than cladogram I, entailing more evolutionary changes and more 
convergences. By the principle of parsimony, cladogram I is preferred 
over cladogram II. 

The stark clarity of cladistics is a recommendation in itself. 
Furthermore, it allows not only the process of cladogram building to be 
computerised but also the ready search for, and objective comparison 
of, alternative cladograms. Cladistics is not, however, completely 
objective and non-arbitrary: the taxonomist’s judgment is important in 
defining characters and judging homology, in particular. ‘The compari- 
son of DNA sequences shows this up most starkly. Because sequences 
evolve both by substitutions of one nucleotide by another and by 
changes in number of nucleotides (by deletion or insertion), there are 
often various ways of explaining differences between homologous 
sequences, depending on the numbers of substitutions and deletions/ 
insertions assumed; parsimony cannot fully resolve such cases because it 
is usually impossible simultaneously to minimise the number of substi- 
tutions assumed and to minimise the number of deletions and insertions. 

Traditional evolutionary taxonomists have a more fundamental 
objection to cladistics than simply that it is not as objective in practice 
as it is in principle. This is that cladograms do not reflect the 
underlying evolutionary tree in the way that the traditionalists would 
like them to do. The problem lies with the cladists’ rejection of 
paraphyletic groups, 1.e. taxa which do not contain all the descendants 


JF. F. D. Greenwood 93 Bull. B.O.C. 1997 117(2) 


of the common ancestor, such as taxon III in Fig. 4. One manifestation 
of the problem arises from traditional classifications attempting to show 
patterns of divergence as well as patterns of branching whereas cladistic 
classifications are concerned only with branching. For example, 
traditionalists separate the Class Aves from the Class Reptilia because 
of the great divergence of birds from reptiles, whereas cladists do not 
allow this because birds are descendants of the common ancestor of 
reptiles (indeed, they are, in branching terms, more closely related to 
crocodiles than either is to turtles or to lizards and snakes). Which 
classification better represents the evolutionary tree depends on one’s 
view of the relative importance of branching and divergence. The 
second manifestation of the paraphyly problem is illustrated by Fig. 5, 
in which species 1 shows the primitive condition for all characters. 
Thus, on the evidence available, the most parsimonious view of the 
evolution of the four species in the Figure is that species 1 is the 
common ancestor of all of the others. However, cladistics involves 
recognising groups by homologies and has no means of distinguishing 
ancestor—descendant relationships. (Cladists would, indeed, argue that 
no-one has such means since, even if species 1 was represented in the 
fossil record at a time compatible with its being the ancestor of the 
others, there is no way of knowing that the fossil specimens were 
certainly ancestral to the other species.) Furthermore, to recognise 
species 1 in Fig. 5 as the ancestor of the other species would make that 
species itself a paraphyletic taxon, since it does not contain all its 
descendants. A practical example, if one assumes Archaeopteryx to 
represent the ancestor of all later birds, is that Archaeopteryx cannot be 
recognised (in a cladistic classification) as a genus equivalent to other 
avian genera, because it would then be a paraphyletic taxon. Cladists 
have attempted to resolve this problem in various ways, though none of 
the solutions produce classifications that non-specialists find easy to 
understand. Whether the matter will be resolved by non-specialists 
becoming sufficiently educated to be able to understand such 
classifications or by the partial acceptance of paraphyletic taxa, I am 
reluctant to predict. 


Pattern cladistics 


Pattern cladistics uses cladistic methods to produce classifications but it 
avoids the uncertainties involved in working out phylogenetic trees and 
the problems that arise if one tries to describe trees through 
classifications by ignoring evolution (for the purposes of classification). 
In brief, it concentrates on the pattern of organic diversity rather than 
on the process that produces the pattern. Ridley (1986) has argued that, 
once evolution is abandoned, there is no justification for cladism (it is 
reduced to just one arbitrary method among many for pigeon-holing 
animals and plants) but pattern cladists would reply that, to study 
evolution, one should describe the patterns first (without preconcep- 
tions) and then worry about the processes. They remain the major force 
in modern taxonomy and their methods have undoubtedly been useful 
in many systematic and biogeographical studies. 


FJ. F. D. Greenwood 94 Bull. B.O.C. 1997 117(2) 


Topics addressed in the symposium 


Endler (1989), in a balanced and illuminating review, has pointed out 
that not only is there a variety of different species concepts but that 
there are several major differences in the aims of species concepts and 
that different concepts have different uses. Much attention at the 
symposium, led particularly by Liversidge, by Zink and by Snow, 
focused on the relative merits of various species concepts, especially the 
biological and the phylogenetic species concepts. The latter, though it 
comes in various forms (see Cracraft 1983, 1989, McKitrick & Zink 
1988, Nelson 1989), can be seen as a logical extension of cladistics to 
the species level, with its concentration on pattern rather than process. 
There is no doubt that, because of uncertainties associated with 
assessing the ‘process’ (potential interbreeding), defining biological 
species is not always easy or objective. Indeed, most evolutionary 
biologists would agree with Templeton (1989) that his ‘“‘cohesion 
species concept’ is generally more useful than the traditional biological 
(“isolation’’) species concept; but this still emphasises process rather 
than pattern. The phylogenetic concept may appear to resolve 
problems by concentrating on pattern but some of us remain to be 
convinced that it will prove more workable in practice. Indeed, 
disputes over trinomials (discussed at the symposium by Violani & 
Barbagli) are symptomatic of the difficulties of defining taxa at levels 
below that of the biological species. ‘The well-known difficulties for the 
biological species concept of deciding whether or not to treat allopatric 
forms as separate species are paralleled for the phylogenetic species 
concept: if one applies the usual criterion under this concept that the 
two populations are to be specifically separated if they are diagnosably 
distinct, then one would separate two populations that differed at only 
a single genetic locus provided that difference was consistent; this 
would not generally be helpful, but the alternative is to impose an 
arbitrary rule about how much difference is required before one treats 
two forms as separate species—just as when applying the biological 
species concept to allopatric forms. 

Which species concept one uses clearly has implications beyond 
systematics (some being explored at the symposium by Knox and by 
Collar) but in my view the critical points are ones that centre on 
systematics itself. Firstly, even though the process of speciation is 
usually gradual and not always a simple branching, the stage at which 
the genetic and ecological cohesion of a species breaks down to produce 
two or more separate daughter species (themselves internally cohesive) 
represents an important discontinuity; the separation itself favours 
more rapid genetic and ecological divergence, so there is a positive 
feedback. Secondly, the criterion of reproductive and ecological 
cohesion results in biological species having an objective existence in a 
way that other taxonomic levels do not—‘“‘the species is not an 
invention of taxonomists or philosophers, but it has a reality in nature”’ 
(Mayr 1988). That is why most modern philosophers of biology reject 
the idea that species are classes, though they may differ as to whether 
they are therefore to be described as individuals (see discussions in 


F. F. D. Greenwood 95 Bull. B.O.C. 1997 117(2) 


Mayr 1988, Hoffman 1989, Sober 1993). Admittedly, this argument 
falls down if, like some cladists (Nelson 1989), one does not believe in 
species. Such extreme views should not lead us to conclude that the 
ideas involved in the phylogenetic species concept have nothing to offer 
in terms of improving our classifications. Equally, the occasional 
difficulties of applying the biological species concept should not lead us 
to abandon it, given its proven value in ornithology over the last half 
century. 

The symposium was not only concerned with taxonomic principles 
and the consequences of applying different species concepts. 
Mikhailov’s presentation on egg-shell structures reminded us of the 
constant search for new characters needed to resolve taxonomic 
problems. We were reminded of the intimate practical connection 
between classification and identification by Parkin’s contribution on 
DNA-based methods for identifying individuals and their relationships 
and by Russell’s presentation on identifying mummified falcons, where 
the nature of the material required the use of characters other than 
those normally employed. 

Taxonomy is not just a subject for the specialists but a subject of 
importance for all biologists. The level of attendance and liveliness 
of debate at the symposium confirm ornithologists’ current interests 
in avian taxonomy and the freshness of the presentations promise 
continuing developments of both ideas and methods. 


Acknowledgements 


I thank Professor C. J. Humphries for most valuable advice on modern taxonomic 
literature and him and Dr A. Knox for their forthright but constructive comments on an 
early draft. 


References: 

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Cain, A. J. 1959. The post-Linnaean development of taxonomy. Proc. Linn. Soc. London 
170: 234-244. 

Cracraft, J. 1983. Species concepts and speciation analysis. Current Ornithology 1: 
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Cracraft, J. 1989. Speciation and its ontology: the empirical consequences of alternative 
species concepts for understanding patterns and processes of differentiation. 
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Sunderland, Mass. 

Cronquist, A. 1988. A botanical critique of cladism. Botanical Review 53: 1-52. 

Dobzhansky, T. 1973. Nothing in biology makes sense except in the light of evolution. 
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Endler, J. A. 1989. Conceptual and other problems in speciation. Pp. 625-648 in D. Otte 
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Forey, P. L., Humphries, C. J., Kitching, I. J., Scotland, R. W., Siebert, D. J. & 
Williams, D. M. 1992. Cladistics: a Practical Course in Systematics. Clarendon Press, 
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Harshman, J. 1994. Reweaving the tapestry: what can we learn from Sibley and Ahlquist 
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Hennig, W. 1950. Grundziige einer Theorie der phylogenetischen Systematik. Deutsche 
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Hennig, W. 1966. Phylogenetic Systematics. Univ. Illinois Press. 


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Jeffrey, C. 1977. Biological Nomenclature. 2nd edn. Edward Arnold, London. 

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Linnaeus, C. 1758. Systema Naturae, Part I. 10th edn. Stockholm. 

Mayr, E. 1969. Principles of Systematic Zoology. McGraw Hill, New York. 

Mayr, E. 1988. Toward a New Philosophy of Biology. Belknap Press, Harvard University. 

McKitrick, M. C. & Zink, R. M. 1988. Species concepts in ornithology. Condor 90: 1-14. 

Nelson, G. J. 1989. Species and taxa: systematics and evolution. Pp. 60-81 in D. Otte & 
J. A. Endler (eds), Speciation and 1ts Consequences. Sinauer, Sunderland, Mass. 

Nuttall, G. H. F. 1904. Blood Immunity and Blood Relationship. Cambridge Univ. Press. 

Patterson, C., Williams, D. M. & Humphries, C. J. 1993. Congruences between 
molecular and morphological phylogenies. Ann. Rev. Ecol. Syst. 24: 153-188. 

Ridley, M. 1986. Evolution and Classification: the Reformation of Cladism. Longman, 
London. 

Sheldon, F. H. & Bledsoe, A. H. 1993. Avian molecular systematics, 1970s to 1990s. Ann. 
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Sibley, C. G. & Ahlquist, J. E. 1990. Phylogeny and Classification of Birds. Yale Univ. 
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Simpson, G. G. 1961. Principles of Animal Taxonomy. Columbia Univ. Press. 

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Address: Dr J. J. D. Greenwood, British Trust for Ornithology, The National Centre for 
Ornithology, The Nunnery, Thetford, Norfolk, [P24 2PU, U.K. 


© British Ornithologists’ Club 1997 


R. M. Zink 97 Bull. B.O.C. 1997 1147(2) 
Species concepts 


by Robert M. Zink 


It is difficult to imagine a concept that impinges on more biological 
research than that of the species. Most biological studies refer to their 
subjects as members of some species (Hauser 1987). We are often 
taught that species are the only units in the classificatory scheme that 
exist independent of taxonomists; i.e. species are real. Given the 
importance of species in theories about evolution, ecology, and 
behaviour, faunal lists, and for communication of our understanding:of 
biodiversity, lack of agreement about how biologists define species is 
surprising. 

The species debate was evident in Darwin’s time (1859), and has 
escaped general resolution in the last 130 years, except perhaps for the 
fact that most agree that the word species derives from Latin meaning 
‘appearance’ with a secondary meaning ‘kind’. Today the literature is 
replete with different definitions (Table 1). Some (e.g. Endler 1989) 
suggest that different species concepts are needed to study different 
evolutionary processes. Paleontologists must cope with incomplete 
fossil histories and absence of information on mating tendencies (Wiley 
1978). Botanists must deal with reticulation, recognizing that a large 
percentage of all plant species, perhaps 50% or more, is of hybrid origin 
(Cronquist 1978). A large number of biologists claim to follow the 
so-called biological species concept (BSC; Mayr 1942), in which 
reproductive isolation is viewed as the crux of speciation. In the past 15 
years, those coming from a background in phylogenetic systematics, or 
‘cladistics’, have made substantial contributions to the debate about 
species, and have penned a number of ‘phylogenetic’ species concepts 
(Cracraft 1983, de Queiroz & Donoghue 1988, Nixon & Wheeler 1990). 
Although argument over species concepts has persisted for more than 
100 years, it is my thesis in this paper that the contributions from 
phylogenetic systematics have substantively changed the nature of the 
debate. Furthermore, it is my opinion that the long-entrenched 
biological species concept is losing favour in ornithology (Zink & 
MeKitrick 1995, Hazevoet 1996) and elsewhere (Mallet 1995). 

Comparison of the species concepts given in Table 1 is beyond the 
scope of this review. Instead, I will contrast the biological species 
concept and a phylogenetic alternative. The reason for doing so is that 
I believe that the current debate about species concepts has come to 
focus on whether to give primary emphasis to a process, such as mate 
choice, or to correct representation of evolutionary patterns, such as 
those recovered by direct analysis of characters (Graybeal 1995). These 
two conceptual positions are embodied in the BSC and a PSC, 
respectively. Here I review each concept, noting some perceived 
strengths and weaknesses (Tables 2, 3). 

The biological species concept has a long history in ornithology 
(Haffer 1992, Amadon & Short 1992, Bock 1986, Hauser 1987). A 
perceived advantage of this concept is that it is ‘biological’. By 


R. M. Zink 98 Bull. B.O.C. 1997, T#7@) 
TABLE 1 


Some species definitions or views on species 


‘““No one definition has as yet satisfied all naturalists; yet every naturalist knows vaguely 
what he means when he speaks of a species.’ (Darwin 1859) 


*“A species is a set of populations capable of combining with each other but not with other 
similar sets of populations on the basis of affinity and co-direction in ecological 
specialization.” (Shaposhnikov 1966) 


‘A species is a group of organisms not itself divisible by phenetic gaps resulting from 
concordant differences in character states (except for morphs such as those resulting from 
sex, caste, or age differences), but separated by such phenetic gaps from other such 
groups.”’ (Michener 1970) 


“‘We may regard as a species (a) the smallest (most homogeneous) cluster that can be 
recognized upon some given criterion as being distinct from other such clusters, or (b) a 
phenetic group of a diversity somewhat below the subgenus category, whether or not it 
contains distinct subclusters.”’ (Sneath & Sokal 1973) 


““Somit ist die Art als das Kollektiv von Lebewesen zu bestimmen, das gemeinsam eine 
dkologische Nische behauptet.’’ (von Wahlert 1973) 


“Species may then be defined as groups of phenetically similar populations that have the 
capability to interbreed, and share similar ecological characteristics.’’ (Doyen & 
Slobodchikoff 1974) 


“Species, then, are the most extensive units in the natural economy such that 
reproductive competition occurs among their parts.’’ (Ghiselin 1975) 


‘A species is a lineage (or a closely related set of lineages) which occupies an adaptive 
zone minimally different from any other lineage in its range and which evolves separately 
from all lineages outside its range.’’ (Van Valen 1976) 


‘Species are the smallest groups that are consistently and persistently distinct, and 
distinguishable by ordinary means.”’ (Cronquist 1978) 


‘““A species is a single lineage of ancestral descendant populations of organisms which 
maintains its identity from other such lineages and which has its own evolutionary 
tendencies and historical fate.’’ (Wiley 1978) 


‘““A ‘species’ is merely ‘a population or group of populations defined by one or more 
apomorphous features, it is also the smallest natural aggregation of individuals with a 
specifiable geographic integrity that can be defined by any current set of analytical 
techniques.” (Rosen 1979) 


‘A species is a group of animals or plants all of which are similar enough in form to be 
considered as minor variations of the same organism. Members of the group normally 


interbreed and reproduce their own kind over considerable periods of time.” (Trueman 
1979) 


‘‘A species is a diagnosable cluster of individuals within which there is a parental pattern 
of ancestry and descent, beyond which there is not, and which exhibits a pattern of 
phylogenetic ancestry and descent among units of like kind.’ (Eldredge & Cracraft 1980) 


‘“‘Species are simply the smallest detected samples of self-perpetuating organisms that 
have unique sets of characters.” (Nelson & Platnick 1981) 


‘‘Each species is an internally similar part of a phylogenetic tree.’> (Willis 1981) 


‘‘We can, therefore, regard as a species that most inclusive population of individual 
biparental organisms which share a common fertilization system.”’ (Paterson 1985) 


‘““An ‘evolutionary species’ is a single lineage of ancestor—descendant populations which 
maintains its identity from other such lineages and which has its own evolutionary 
tendencies and historical fate.’’ (Wiley 1978) 


‘fA species is what a good taxonomist says it is.”’ (Anon.) 


R. M. Zink 99 Bull. B.O.C. 1997 117(2) 
TABLE 1 continued 


“At the outset I confess a disbelief in species, as that word is commonly understood to 
refer to the basic taxonomic unit or to the taxonomic unit of evolution .. . There seem to 
be no basic taxonomic units and no particular taxonomic unit of evolution ... and as 
Agassiz said in 1859 ‘species do not exist in nature in a different way from the higher 
groups’.”’ (Nelson 1989) 


a9 


. species as the most inclusive group of organisms having the potential for genetic 
and/or demographic exchangeability.’’ (Templeton 1989) 


A species is “the smallest aggregation of populations (sexual) or lineages (asexual) 
diagnosable by a unique combination of character states in comparable individuals 
(semaphoronts)’’. (Nixon & Wheeler 1990) 


c 


Species “‘refer to groups of actually or potentially interbreeding populations isolated by 
intrinsic reproductive barriers from other such groups. Evidence for reproductive 
barriers . . . will involve concordant genetic differences among the populations involved’’. 
““Subspecies are groups of actually or potentially interbreeding populations phylogeneti- 
cally distinguishable from, but reproductively compatible with, other such groups. 
Importantly, the evidence for phylogenetic distinction must normally come from the 
concordant distributions of multiple, independent, genetically based traits.” (Avise & 
Ball 1990) 


“Species are groups of actually or potentially interbreeding natural populations, which 
are reproductively isolated from other such groups.’’ (Mayr 1942) 


“A species is a reproductive community of populations (reproductively isolated from 
others) that occupies a specific niche in nature.’’ (Mayr 1982) 


““A species is the smallest diagnosable cluster of individual organisms within which there 
is a parental pattern of ancestry and descent.” (Cracraft 1983) 


Species are “lineages whose components (if distinguishable) are not incontrovertibly on 
different phylogenetic trajectories (i.e. sublineages, if distinguishable, are reproductively 
compatible), as long as these sublineages do not form a paraphyletic group in recovered 
history’. ““The species category ... would represent the largest entities that have evolved 
whose parts, if distinguishable, are not likely to be on different phylogenetic trajectories.” 
(Frost & Hillis 1990) 


“if a given historical group of hybridogens is persistent and is not affecting the 
evolutionary trajectory of its Mendelian ancestor (as indicated by biogeography, habitat 
preferences, or genetic divergence), it should be considered a separate species.” (Echelle 
1990) 


A species is the “‘smallest recognizable monophyletic or unresolved unit’’. (Donoghue 


1985) 


“Phylogenetic species can be delimited by a procedure (population aggregation analysis) 
that involves a search for fixed differences among local populations, followed by 
successive rounds of aggregation of populations and previously aggregated population 
groups that are not distinct from each other.’ “descent relationships among 
[phylogenetic species] must be hierarchic.’ (Davis & Nixon 1992) 


observing birds from differentiated groups that interact in sympatry, 
one can determine if they mate assortatively. Because we observe the 
birds themselves choosing mates, this is deemed ‘biological’. However, 
there is nothing ‘non-biological’ about the evolution of groups of 
individuals, the hallmark of the PSC, and the BSC cannot make a claim 
to be uniquely biological. Perhaps the ‘B’ in ‘BSC’ should stand for 
behavioural. 


R. M. Zink 100 Bull. B.O.C. 1997 117(2) 


TABLE 2 
Perceived strengths and weaknesses of the biological species concept (see Cracraft 1983, 
McKitrick & Zink 1988, Zink & McKitrick 1995, Zink 1996b) 


Perceived strengths 
Reproductive isolating mechanisms objective in sympatry 
Reproductive isolation= genetic closure of a lineage 


Perceived weaknesses 
Amount of hybridization required for conspecifity is vague 
Stable hybrid zones are difficult to assess 
Time to fusion is potentially enormous 
Allopatric populations are judged subjectively 
Evidence of evolutionary pattern is “‘overruled”’ by actual or presumed interbreeding 
Non-historical species result because hybridization is not limited to sister taxa 
Reproductive isolation is an epiphenomenon, not a directly measured characteristic of 
allopatric populations 


TABLE 3 
Perceived strengths and weaknesses of a Phylogenetic Species Concept 


Perceived strengths 
Species limits are consistent with recovered phylogenetic patterns 
Same procedures used for species limits as for higher taxonomic categories 
Based directly on character variation, not an epiphenomenon such as mate choice 
Better description of spatial patterns of biodiversity 
Produces units required by evolutionary theories and conservation biology 


Perceived weaknesses 
Lower limits of species require careful study of variation 
Limits of diagnosability problematic 
Too many species will result 
Species might be too ephemeral 


The crux of the BSC is that species are reproductively isolated—they 
have reached that stage in evolutionary divergence where members no 
longer recognize individuals in other species as mates, or if they do, 
their offspring are of reduced fitness. Conversely, taxa that are 
reproductively compatible, especially if they interact in sympatry and 
form a hybrid swarm, are considered to be conspecific. Thus, the 
process of mate choice is accorded primary significance in determining 
whether two taxa are to be considered one or two species. This view has 
intuitive appeal, although I believe that the theoretical and practical 
flaws with the BSC are insurmountable. 

One might argue that in practice the BSC cannot be deemed of 
primary significance because taxonomists working with some of the 
most speciose groups actually do not appear to use it. It would be hard 
to argue (see Whittemore 1993) that botanists use the BSC when 
delimiting species, given the propensity of plants to hybridize. 
Similarly, it is doubtful if entomologists (perhaps excluding 


R. M. Zink 101 Bull. B.O.C. 1997 117(2) 


lepidopterists) use the BSC, depending instead on morphological 
differences to recognize most species boundaries. If botanists and 
entomologists do not generally apply the BSC when describing species, 
it follows that the bulk of the world’s biodiversity 1s classified primarily 
by a non-BSC paradigm. ; 

Controversy has surrounded the BSC since its inception (Donoghue 
1985, Mallet 1995), although many believe that it has survived the tests 
of time (Coyne ez al. 1988). Many problems with using the BSC are 
familiar (Cracraft 1983), such as the need to speculate whether 
allopatric populations are reproductively isolated (e.g. ‘Thompson 
1991). Zink & McKitrick (1995) reviewed how the significance of 
hybridization has been misconstrued by proponents of the BSC. It has 
been implied that hybridization might ‘erase’ evolved differences 
between two taxa. However, fusion of hybridizing taxa will likely 
require long time periods (Zink & McKitrick 1995). Thus, the BSC has 
been criticized, rightly in my opinion, for basing species limits on what 
might happen in the future rather than what has happened during the 
evolutionary past (Cracraft 1983). 

The primary new criticism of the BSC to emerge from the writings 
of phylogenetic systematists is the recognition that reproductive 
isolation often does not evolve concomitantly with characters that 
delimit evolutionary taxa, especially in the early stages of divergence. 
Within many biological species we can recognize separate evolutionary 
groups, and analyses of their relationships can suggest a pattern that is 
inconsistent with the pattern of reproductive compatibility—i.e., two 
hybridizing taxa might not be each other’s nearest relatives. A ‘species’ 
consisting of two or more groups that are not each other’s nearest 
relatives is unacceptable to modern systematists. In systematics terms, 
reproductive compatibility is an ancestral condition, because individ- 
uals in the ancestral population recognized each other as mates (Rosen 
1979). To use the primitive ability to hybridize (perceived in allopatry 
or documented in sympatry) as a grouping criterion for species limits, 
as the BSC requires, violates the way in which systematists unite taxa 
in an evolutionary manner—by possession of shared-derived (i.e. 
non-ancestral) characters (Rosen 1979). Species limits based on 
reproductive compatibility (BSC) or patterns of character variation 
(PSC) can conflict and lead to different species limits from the same 
data set (Frost & Hillis 1990) (see Fox Sparrow Passerella iliaca 
example below). 

The above comments do not mean that advocates of a PSC consider 
reproductive isolation uninteresting or unworthy of study. Indeed, one 
could argue that without reproductive isolating mechanisms, the 
world’s species diversity would be kept low because taxa could not 
become sympatric. Reproductive isolation is an inevitable but 
temporally unpredictable by-product of evolutionary divergence 
(McKitrick & Zink 1988). At some point in evolutionary divergence, 
nearly all differentiated taxa are reproductively isolated, and all 
reproductively isolated taxa are likely to be ‘good’ phylogenetic species. 
I suspect that all species concepts recognize reproductively isolated taxa 
as different species. However, between the time of the emergence of 


R. M. Zink 102 Bull. B.O.C. 1997 117(2) 


taxa on their own evolutionary trajectories (as evidenced by characters) 
and their eventual reproductive isolation, the pattern of reproductive 
compatibility is an unreliable predictor of historical relationships of 
taxa. Although recognizing the intrinsic importance of reproductive 
isolation, users of a PSC choose not to include this information in the 
delimitation of species (Frost & Hillis 1990). In fact, proper study of 
the evolution of an attribute such as reproductive isolation requires first 
that historical patterns among taxa are known (Brooks & McLennan 
1990). The potential for non-historical groupings together with the 
other often-noted problems (e.g. Donoghue 1985) leads to the in- 
evitable conclusion that the BSC should be replaced with a concept that 
correctly represents history. 

Debate continues over how to use information on evolutionary 
pattern to delimit species. For example, several phylogenetic species 
concepts exist (Table 1). Davis & Nixon (1992) suggest that the phrase 
“phylogenetic species concept” is misleading because the point is to 
delimit terminal taxa for phylogenetic analysis, and they describe a 
process whereby phylogenetic analysis does not play a role in species 
delimitation. Rather, they show how morphological or molecular 
character evidence can be used alone to delimit species. Unlike the 
BSC, characters are not weighted by their presumed role in a process 
such as mate choice. Nonetheless, common to history-based concepts, 
including the evolutionary species concept (Wiley 1978), is the (1) 
rejection of reproductive compatibility as the primary criterion of 
conspecific status, and (2) recognition that species can hybridize owing 
to the retention of the ancestral ability to do so. In my opinion, the crux 
of a phylogenetic species concept is to recognize groups of individuals ~ 
that have been on independent evolutionary trajectories. Evaluation of 
multiple characters does not allow further subdivision of such groups. 
That is, a PSC attempts to recognize the status guo—character analysis 
reveals groups of individuals that qualify as basal evolutionary units 
(Cracraft 1983, 1989). Nonetheless, ongoing debate revolves around 
how best to recognize history at the population level (Davis & Nixon 
1992, Zink & McKitrick 1995). 

Several criticisms of phylogenetic species concepts exist. Avise & Ball 
(1990) suggested that with modern molecular methods, each individual 
would be diagnosable, and might qualify as a separate phylogenetic 
species. Indeed, if one studied individual gene genealogies (i.e. single 
characters), one would likely be able to circumscribe groups. of 
individuals on a gene-by-gene basis that are not mutually exclusive, 
historical entities (see Maddison 1995). One might then think they are 
forced to recognize individual organisms as phylogenetic species to 
escape the problem raised by conflicting gene genealogies. However, 
the opposite trend is actually true. One uses multiple (unweighted) 
characters and resolves conflicts by an explicit a prior criterion (e.g. 
Davis & Nixon 1992). Species limits are set where character evidence 
becomes maximally congruent. Some character conflicts might remain, 
as they do in most phylogenetic studies. Thus, rather than species 
being single individuals, species often will become geographically 
coherent groupings of individuals. Resolution of conflicting character 


R. M. Zink 103 Ball. B.O.C. 997 P72) 


i P. iliaca 


P. schistacea 


P. megarhyncha * 


\ 


Figure 1. Approximate breeding distribution of four phylogenetic species of the Fox 
Sparrow Passerella iliaca. Phylogenetic tree derived from pattern of restriction sites (Zink 
1994). 


distributions (e.g. gene genealogies) results in more, not less, inclusive 
groupings of individuals. The mistaken belief that advocates of a 
phylogenetic species concept rely on single characters to delimit species 
has misled several authors (e.g. Amadon & Short 1992). Put another 
way, a problem with avian subspecies is that they are often based on 
single characters, and analysis of other characters can suggest different 
subspecific limits (see below). No taxonomic category, species or 
otherwise, will likely be based on single characters (Barrowclough 
1982). 

Differences between biological and phylogenetic species concepts are 
illustrated by studies of the Fox Sparrow Passerella iliaca. In North 
America, Fox Sparrows range over the taiga, northwest coast, and 
mountainous regions of the west (Fig. 1). There is considerable 
phenotypic variation, partitioned by taxonomists into 18 subspecies. 
However, four basic groups exist: zliaca, unalaschcensis, megarhyncha, 
and schistacea. Each of the four groups was originally recognized as a 
separate species. The Fox Sparrow is currently considered a single 
biological species because each group is known to hybridize with at 
least one other group, although only a few hybrid specimens between 


R. M. Zink 104 Bull. B.O.C. 099d TT) 


tliaca and unalaschcensis are known (Zink & McKitrick 1995). Zink 
(1994) found that each of the four groups possessed a distinct set of 
mitochondrial DNA haplotypes. Thus, both genetic and morphological 
evidence reveal four groups of individuals with separate evolutionary 
histories—hence, four likely phylogenetic species. (Note that molecular 
analysis did not indicate a greater number of taxa than that apparent 
from morphological comparisons.) Zink (1994) suggested, however, that 
further study of morphology was needed to test and refine species limits. 

The question of how many biological species of Fox Sparrow exist 
depends on how one interprets the evidence on hybridization. An 
extreme view supports one species, because at least a trickle of genes 
links all parapatric forms. A moderate view might envisage three 
species, zliaca, unalaschcensis, and (megarhyncha plus schistacea). The 
latter grouping would obtain because of a narrow hybrid zone between 
the latter two taxa (Zink 1994). 

The main problem in interpreting biological species limits in the Fox 
Sparrow concerns megarhyncha and _ schistacea. MtDNA evidence 
reveals a narrow hybrid zone between the two groups (Zink 1994). The 
zone seems broader when morphometric patterns are considered. The 
stability of the zone is unknown, a factor critical in interpreting 
biological species limits (Mayr 1982, Zink & McKitrick 1995). If the 
hybrid zone was stable, Mayr (1982) would consider schistacea and 
megarhyncha to represent two species because there was an unknown 
barrier to complete introgression despite random mating in the zone. 
Because the zone appears to be between two non-sister taxa, it is 
probably one of secondary contact (Cracraft 1989) between two 
phylogenetic species that have retained the primitive ability to 
hybridize. This study therefore illustrates the problem identified with 
the BSC by phylogenetic systematists (e.g. Rosen 1979); other avian 
examples are found in Moore et al. (1991) and Freeman & Zink (1995). 
Lumping schistacea and megarhyncha into a single species would 
mis-represent evolutionary history, because although they hybridize 
extensively, they are not each other’s nearest relatives (Fig. 1). It could 
take tens of thousands of generations for significant introgression to 
occur (Zink & McKitrick 1995). The PSC would recognize the status 
quo, namely that these two taxa have had independent evolutionary 
histories and are therefore (phylogenetic) species. Biological species 
limits depend on one’s interpretation of how much (or little) 
hybridization is required. BSC advocates could therefore recognize 1, 
2, 3, or 4 species (which should challenge the belief of those who think 
that species are real entities of nature). However, a non-historical 
species including only megarhyncha and schistacea, permissible under 
the BSC, would be of no intrinsic value in phylogeny studies, speciation 
research, biogeography, comparative biology or conservation. I can 
think of few studies that would intentionally use this classification. 
Non-historical taxa, then, are the Achilles heel of the BSC. 

It is worth noting that the inconsistent relationship between patterns 
of evolution among populations and their reproductive isolation has 
only recently been recognized, in part owing to molecular systematics. 
Molecular methods, such as DNA sequencing, can resolve evolutionary 


R. M. Zink 105 Bull. B.O.C. 1997 117(2) 


relationships over short evolutionary time scales, such as within 
biological species. Prior to molecular analysis, relatively little attention 
focused on phylogenetic relationships among, for example, subspecies 
(including those in the Fox Sparrow). Molecular studies can, of course, 
reveal taxa ‘invisible’ to morphologists, but more often they elucidate 
inter-relationships of closely related taxa. Thus, the BSC was invented 
and used commonly before it was realized that hybridizing taxa might 
not be sister taxa, and that the pattern of hybridization might 
misrepresent the true pattern of evolutionary history. Now, however, 
molecular phylogenetic analysis and classification (i.e. cladistic) 
methods that require taxonomic boundaries to be faithful to evol- 
utionary ones, mean that the BSC is not an appropriate species concept. 

Given the historical usage of the BSC in ornithology, one might 
wonder if it can be salvaged. The BSC would be improved if subspecies 
were phylogenetic species that were not reproductively isolated. 
Species would be required to be consistent with recovered patterns of 
history (thus, the ‘Fox Sparrow’ would be either one, two or four 
species, but one would not accept a species consisting solely of 
megarhyncha and schistacea given the pattern of history suggested in 
Fig. 1). Notwithstanding potential improvements to the BSC, ‘species’ 
would still contain variable numbers of basal evolutionary taxa, and 
thus not be comparable. Also species membership of allopatric 
subspecies would be judged subjectively, and still one would have to 
guess whether an allopatric taxon was reproductively isolated rather 
than emphasizing directly observable character variation. I think that 
scientific studies require more of species concepts than can be 
accomplished by re-modelling the BSC. 

Implementation of a phylogenetic species concept would remove 
another contentious area from avian systematics, the subspecies 
category. Many practising avian taxonomists have in mind a certain 
‘level of distinctness’ required to elevate a subspecies to species; 
unfortunately, this level varies from taxonomist to taxonomist because 
of subjective character weighting. Under a PSC, there is no need for 
the subspecies category (McKitrick & Zink 1988). I suggest that this 
would be beneficial because of the many avian subspecies that represent 
arbitrary divisions of clines, or are based on characters not supported 
by, or conflicting with, other characters. Thus, there would be a single 
taxonomic category for basal evolutionary taxa—phylogenetic species. 

Mayr (1993) was concerned that implementation of a PSC would 
double the number of biological species of birds (c. 9000) recognized 
worldwide. Mayr implied that this number of species would hinder 
ornithological research. If this were so, botanists and entomologists 
surely would be unable to conduct research. G. F. Barrowclough, 
J. Cracraft & R. M. Zink (unpubl. data) estimated that c. 18,000 species 
of birds exist, using the PSC. This estimate was reached by evaluating 
morphological evidence that was gathered by previous avian 
taxonomists for subspecies designations; in fact, many subspecies did 
not qualify as phylogenetic species (see Hazevoet 1996). This number 
of species can hardly be perceived as a negative aspect of a PSC 
(Cracraft 1992). I suggest that it would (1) more accurately portray 


R. M. Zink 106 Bull. B.O.C. \997 17 


avian species diversity (Moritz 1994, Rojas 1992, Cracraft 1997), and 
(2) put the assessment of avian biodiversity on a more comparable 
footing with other groups of organisms. 

Martin (1996) suggested that use of molecular methods would 
drastically inflate the number of avian phylogenetic species, many of 
which would be recognizable only with sophisticated molecular tools 
(i.e. beyond field identification). Zink (1996a) suggested that the 
number of new species that were diagnosable only through molecular 
analyses would in fact be relatively low, and that most taxa supported 
by molecular analyses also have morphological characteristics (Zink 
et al. 1995). The doubling of the number of bird species mentioned 
above (G. F. Barrowclough et al., unpubl. data) was based solely on 
morphological criteria. At the Linnean symposium from which this 
paper derives, two independent speakers estimated the number of 
phylogenetic species for particular areas by examination of published 
(morphological) taxonomies. Thus, I doubt that molecular analyses will 
reveal many species that are ‘invisible’ to field workers (see the Fox 
Sparrow example above), and even if they did, would we recognize as 
valid components of our biodiversity only those taxa that could be 
identified with binoculars and a certain level of expertise? 

A phylogenetic species concept would also benefit conservation 
biology (Cracraft 1997). Today, under the BSC, one needs to argue that 
sometimes local populations, subspecies, or species are units that 
should be targeted for conservation efforts. In my opinion, many non- 
systematists (and systematists) view this inconsistency as unscientific. 
Under a phylogenetic species concept, phylogenetic species would 
become the category for conservation biology. The explicit goal would 
be to preserve biodiversity at its most basic spatial scale. It might be 
thought that some phylogenetic species, formerly ‘only’ subspecies, 
would be indefensible for conservation efforts. Because conservation 
efforts are likely to move from the level of individual species to 
communities, patterns of species diversity will be used to define 
conservation entities, such as reserves that capture the bulk of areas of 
endemism. Spatial patterns of species diversity are most consistently 
and precisely described under a PSC. 

Is the species debate important? Clearly, if different species concepts 
lead to different species limits given the same data, the answer is yes. 
The species debate is not a semantic battle between cladists and 
evolutionary taxonomists. Many researchers are studying speciation. If 
we cannot agree on a concept of species, how can speciation be 
effectively studied (Zink 1996b)? If we are discussing how to preserve 
biodiversity, and species are our measure, species concepts are 
extremely important. If we use species in evolutionary theories, or 
comparative studies (Brooks & McLennan 1990), we must be in 
agreement about how they are described. We must understand 
properties of species under differing concepts and determine that these 
properties are what our theories and uses of species require. 
Researchers should recognize the impact that different species concepts 
can have on their research, and explicitly consider whether their 
research programme depends on which concept is used. 


R. M. Zink 107 Bull. B.O.C. 1997 117(2) 


Today, the seemingly perennial species debate has been materially 
changed by contributions from phylogenetic systematics, and has come 
to involve whether to represent evolutionary history faithfully by 
species limits. I think that the BSC is theoretically flawed because it can 
lead to, and accept, misleading historical groupings, and because there 
has not been a satisfactory resolution to the problem of classifying 
allopatric populations. Problems with a PSC involve mainly practical 
ones, such as how to delimit basal taxa, an impressive start at which was 
made by Davis & Nixon (1992). Although there will be phylogenetic 
species whose limits are ‘fuzzy’ owing to the dynamic nature of 
evolutionary change, and specific individuals difficult to place into a 
particular species, I think that these problems are far less important 
than the theoretical and practical ones encountered by applying the 
BSC. Giving primacy to correct representation of history should be the 
basis of our species concept, and it will produce species that are best 
(not perfectly) suited to the majority of biological uses to which they 
are put. A species concept consistent with a phylogenetic species 
concept should be adopted in ornithology, replacing the BSC. 


Acknowledgements 


I am grateful to many individuals who have discussed species concepts with me for years: 
G. F. Barrowclough, J. Cracraft, S. J. Hackett, J. M. Bates, J. Davis, M.S. Hafner, J. V. 
Remsen, J. T. Klicka, S. M. Lanyon, E. C. Birney, S. J. Weller, D. P. Pashley and M. C. 
McKitrick; mention in this list does not, however, imply agreement with the views stated 
in this paper. I thank S. M. Lanyon, J. T. Klicka, E. C. Birney and S. J. Weller and two 
anonymous reviewers for comments on the manuscript. Some of Table 1 was contributed 
by D. B. Wake. A. Fry made Figure 1. 


Summary 


The debate over species concepts is in an active phase. After years of acceptance of the 
biological species concept (BSC) in ornithology and other disciplines, the field of 
phylogenetic systematics has contributed a significant new challenge. Reproductive 
isolation, the hallmark of the evolution of biological species, does not necessarily evolve in 
concert with characters that reflect the pattern of population subdivision. That is, taxa on 
independent evolutionary trajectories, only some of which might be reproductively 
isolated, can exist within biological species. Therefore, setting species limits consistent 
with patterns of reproductive compatibility can lead to species limits that misrepresent 
evolutionary history because hybridizing taxa might not be each other’s nearest relatives. 
A phylogenetic species concept (PSC) equates species with groups of evolutionarily 
distinct groups of individuals that cannot be further subdivided by analysis of multiple 
characters, irrespective of mating tendencies. It requires that species limits are consistent 
with known patterns of evolutionary history. The PSC recognizes that (phylogenetic) 
species can hybridize because they retain the primitive ability to do so. The debate over 
species concepts currently focuses on whether to give primary emphasis to reproductive 
isolation and the process of mate choice (BSC), or to historical patterns of character 
variation (PSC); this distinction results in different species concepts leading to different 
species limits given the same data, such as in the example discussed of the Fox Sparrow 
Passerella iliaca. 1 suggest that a version of the PSC should replace the biological species 
concept. This would serve several useful functions, such as (1) making species of birds 
more equivalent with species in other major taxonomic groups, (2) providing an objective 
method for classifying allopatric populations, (3) removing the contentious category of 
subspecies, and (4) ensuring that species limits are consistent with recovered historical 
patterns. Fears that a PSC coupled with molecular methods would produce too many 
species are unfounded. The units required by phylogenetic analyses, comparative 
ethological, evolutionary and ecological studies, biogeography, and conservation biology 
are in practice phylogenetic species; biological species can fail these needs. 


R. M. Zink 108 Bull. B.O.C. 1997 17@) 


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Address: Dr R. M. Zink, J. F. Bell Museum, 100 Ecology Building, University of 
Minnesota, St. Paul, MN 55108, U.S.A. 


© British Ornithologists’ Club 1997 


D. W. Snow 110 Bull. B:O:C. 1997 Wa @ 


Should the biological be superseded by the 
phylogenetic species concept? 


by D. W. Snow 


A controversy of fundamental importance has been developing in 
animal taxonomy in recent years, and is beginning to have practical 
effects on ornithology. It is the argument that the long-accepted 
biological species concept (hereafter BSC)—as expounded notably by 
E. Mayr in influential publications since 1942—1is fundamentally flawed 
and should be replaced by the concept of the ‘phylogenetic species’. 
Hazevoet’s (1995) excellent check-list of the birds of the Cape Verde 
Islands, the latest in the B.O.U. check-list series, adopts the 
phylogenetic species concept (hereafter PSC), and is the first 
authoritative work on an entire, though small, avifauna to do so. 
Cracraft’s (1992) reclassification of the birds-of-paradise was the first, 
and still remains the only, such treatment of a bird family; Livezey 
(1991, 1995a, b) has. applied the PSC to three tribes of the Anatidae. 

The principles underlying the PSC have been expounded by Cracraft 
(e.g. 1983, 1987, 1992), Sluys (1991) and several other authors in the 
U.S.A., where the concept originated; McKitrick & Zink (1988) have 
advocated its use in ornithology. For most British readers the most 
accessible, concise introduction to it will be the clear and forceful 
4-page exposition by Hazevoet in Appendix 4 to his check-list. 

The purpose of the present paper is not to discuss at length the 
arguments for and against different species concepts. This has been 
done in a number of publications, including those mentioned above. Its 
main purpose is to mention briefly what seems a fundamental weakness 
of the PSC; to argue that the flaws in the BSC, while not being denied, 
have been exaggerated; and finally to discuss in greater detail the 
practical difficulties that would be encountered in any attempt to apply 
the PSC wholesale in avian taxonomy. 


The phylogenetic species 


Cracraft (1983) defines a phylogenetic species as “the smallest 
diagnosable cluster of individual organisms within which there is a 
parental pattern of ancestry and descent’’. Other definitions have been 
given, but all agree that the species should be defined as the smallest 
distinct group or population of common (monophyletic) ancestry; 
that individuals within the group must normally interbreed only with 
other members of their group; and that whether or not they are 
reproductively isolated from members of other such groups (in the 
sense of not interbreeding successfully with them in the event of their 
coming into contact) is not relevant to their species status. Species are 
therefore the present end-products of evolutionary change, or 
“evolutionary units’’. 

It is at once obvious that, if the PSC is adopted, there will be a huge 
increase in the number of bird species. The species recognised will 


D. W. Snow ci Ball: B:OsE A995 187 (2) 


comprise every monotypic species and many of the subspecies 
recognised under the biological species concept. The total will increase 
from c. 9000 to perhaps 20,000 (the latter figure based largely on 
guesswork; many of the 27—28,000 subspecies and monotypic species 
estimated by Mayr & Gerloff (1994) would not be recognised as 
phylogenetic species—see below). One must agree with Hazevoet 
(1995), however, that the sheer number of species recognised under any 
species concept should not in itself be a factor of any importance in the 
argument, which is based on principle not convenience. The critical 
issues are two: first, whether the PSC is biologically sounder than the 
BSC; and second, whether its application would be practicable. 


Validity of the biological species concept 


The BSC is now so well entrenched in almost all standard 
ornithological literature that it is taken for granted by all ornithologists 
who are not primarily taxonomists, and by most taxonomists too. But 
there are difficulties in applying it, some of which are in practice 
insuperable. Insuperable practical difficulties do not, of course, prove 
that the concept is unsound. The main criticisms of the BSC— 
defined as “groups of actually or potentially interbreeding natural 
populations which are reproductively isolated from other such groups”’ 
(Mayr 1942), with later variants which are not fundamentally 
different—are threefold: (1) that it is ‘process-based’, that is, based 
on an inferred speciation process, not on a rigorous analysis of 
taxonomically valid characters; (2) that the criterion of reproductive 
isolation is almost always untestable, so that informed guesses have to 
be made about the status of more or less closely related allopatric 
populations; and (3) that the polytypic species recognised under the 
BSC are not single irreducible evolutionary units; some (monotypic 
species) are, others (polytypic species) are not. Other objections that 
have been made seem less crucial, as they result largely from 
misapplication of the BSC through insufficient knowledge or 
inadequate analysis; for instance, that some polytypic species later turn 
out not to be monophyletic (with the likelihood that there are other 
such, not yet detected). 

It cannot be denied that the first two of these major criticisms have 
some validity. The first is valid insofar as it reflects on the way in which 
some bird taxonomists have worked in the last 50 years; but as is argued 
below, cladistic analysis of taxonomic characters would in the great 
majority of cases almost certainly confirm the validity of the 
assemblages recognised by the BSC as polytypic species. The second 
criticism has long been recognised as the main stumbling block 
preventing a consistent and generally accepted application of the BSC. 
Decisions about the specific status of well-marked allopatric forms are 
somewhat arbitrary. They may even seem to be susceptible to changing 
fashions, as shown by the present trend towards giving species status to 
forms that have previously been treated as subspecies. In fact, such 
changes are usually a consequence of fuller knowledge of vocal 
differences and other potential isolating mechanisms; but see Collar 


D. W. Snow 112 Bull. B:O.C, 1997, TV 


1997 (p. 130 in this issue). If the allopatric model of speciation is 
valid—and proponents of the PSC do not argue against it—this 
problem is to be expected. In any process of gradual change from one 
category to another there will be borderline cases; indeed it seems, as 
mentioned below, that the PSC encounters the same problem. 

The third main criticism of the BSC is of a rather different kind. It 
concerns a matter of definition, and in my view is a main weakness of 
the PSC. It is not at once obvious why it is right to define species in the 
way the PSC defines them, rather than in the way the BSC does. Why, 
in fact, should all the distinct, even if only slightly distinct, ‘end twigs’ 
of the avian family tree be ranked as species? I know of no convincing 
answer. ‘The idea seems to have a quasi-philosophical rather than a 
biological basis, and tends to be presented as dogma. Thus Cracraft 
(1992) writes: “The proper taxonomic framework for counting 
biological diversity resides with taxa of species rank, not with 
subspecies as required by the biological species concept.’ One is at 
liberty to disagree, and to answer that it is equally proper to use 
subspecies (as well, of course, as monotypic species). I suggest that this 
is a matter of opinion, not of science. 

Defence of the BSC, at least in birds, can be argued more positively. 
Surely any ornithologist who has studied any group of birds with a 
wide geographical range (except perhaps some oceanic birds) must be 
convinced of the reality of what we have become accustomed to call 
‘polytypic species’, however the units comprising them may be 
designated by taxonomists. The facts are so well known that it may 
seem unnecessary to labour them, but in the present context it is 
necessary. ‘Typical Blackbirds Turdus merula—differing slightly in 
colour, but more in size and proportions; some very large, some very 
small, some of intermediate size; with rather different wing-shapes 
depending on whether they are migratory, resident, confined to oceanic 
islands, etc.—comprise an assemblage of forms which not only differ 
from other such assemblages but only make evolutionary sense on the 
assumption that they have been derived from an ancestral form 
probably centred on the west Palaearctic. This is an inference arrived 
at, not perhaps by cladistic analysis of all valid taxonomic characters 
(though such an analysis would almost certainly support the relatedness 
of the members of the assemblage), but by that generally very reliable 
computer, the unaided human brain. The fact that there are other more 
distinct, allopatric merula-like forms in the east Palaearctic and Oriental 
regions (maximus of the Himalayas, mandarinus of China, the simillimus 
group of the Indian subcontinent), and that it is impossible with 
present knowledge to know which, if any, of them might prove to be 
reproductively isolated, i.e. behave as separate species, from west 
Palaearctic merula, may be inconvenient for those wanting a stable 
taxonomy but is entirely consistent with the allopatric (Mayrian) model 
of speciation. The difficulty of deciding the taxonomic status of 
merula-like birds in the east is reasonably explained by their longer 
geographical isolation from western Blackbirds, and the very different 
environmental influences to which they have been subject and become 
adapted. 


D. W. Snow 113 Bull. B.O.C. 1997 117(2) 


This kind of pattern of variation is all-pervasive; its details are 
summarised in regional handbooks, and dealt with in greater detail in 
monographs and other specialised publications. Attention is usually 
drawn to cases in which there is doubt as to the best taxonomic 
treatment under the BSC. Revisions are often made, as further data 
become available, but the main framework, based on the polytypic 
species, remains intact. It is reasonable to argue that the onus is on the 
advocates of the PSC to show, not only that the BSC is not a 
satisfactory framework for the classification of the different kinds of 
birds existing in the world today, but also that adoption of the PSC 
would be fraught with fewer problems. 


Problems of the PSC 


Cracraft’s (1983) definition of a species has been quoted above. Nelson 
& Platnick (1981) give a very similar definition: “Species are simply 
the smallest detected samples [=populations, presumably] of self- 
perpetuating organisms which have unique sets of characters’’. In both 
definitions, the essential points are that they are the smallest groups of 
individuals and that their set of characters is diagnosable as unique. 
This seems simple enough, but the application of the definition would 
in very many cases be by no means easy. In the first place, the samples 
available are often not very large; in birds, their effective size would be 
further reduced by the need to compare specimens of the same age and 
sex class, and with plumage in a comparable state (breeding or 
non-breeding, degree of wear, etc.). Size differences may often be 
critical; but in passerines, for example, first-year birds are usually a 
little shorter-winged than older birds, and it is not always easy to 
distinguish them by plumage. Slight colour differences, which may be 
diagnostic in fresh plumage, may not be apparent in worn plumage. For 
the BSC, such sources of variation may make it hard to assess 
subspecies, but it is well understood that there may be differences of 
opinion about subspecies; it is not very important. For the PSC, where 
the decision affects the recognition or non-recognition of a species, the 
problem is acute. Two examples from the west Palaearctic may 
illustrate the kind of difficulty that will make application of the PSC 
hard to achieve with any hope of a consensus or of stability. 

The kind of geographical variation shown by Turdus merula has 
already been mentioned. At the two extremes of size (based on male 
wing-length) are JT. m. maximus of the Himalayas (144-167 mm) and 
T. m. azorensis from the Azores (117-127 mm). T. m. maximus should 
perhaps be treated as specifically distinct from the west Palaearctic 
forms under either of the debated species concepts; not only is it huge, 
but it differs quite markedly in plumage, and also in song and calls 
(Roberts 1992). But leaving aside maximus, T. m. intermedius of the 
central Asian mountains, which has typical merula plumage, is also very 
large, with a long bill, its wing-length (130-143 mm) falling outside the 
range of azorensis. T. m. syriacus, of Turkey and the Levant, may form 
a link with the European populations, being intermediate in size, but 
further study of these central and southwest Asian populations would 


D. W. Snow 114 Bull. B.O.C. 1997 117(2) 


be needed in order to establish how isolated they are from one another 
and whether variation is clinal. At the bottom end of the size range, 
T. m. cabrerae of Madeira and the Canary Islands (122-129 mm) is 
about as small as azorensis; it differs form azorensis in having darker 
female plumage, a less rounded wing, and a slightly longer tail. There is 
some variation within the Canary Islands, which needs further study. 
In Europe and North Africa, between the very large eastern continental 
populations and the very small Atlantic island populations, there is 
clinal variation which the BSC recognises by a number of subspecific 
names. It is generally admitted that their boundaries cannot be exactly 
defined (e.g. Cramp 1988). 

If there is some difficulty under the BSC in deciding whether the 
eastern Palaearctic and Oriental groups of Blackbird-like forms should 
be treated as conspecific with merula, the problem for the PSC would 
involve the whole range, from the Atlantic islands to China. The 
Azores population would certainly be a phylogenetic species; the 
Madeiran and Canary Islands populations would demand detailed 
study. hey are certainly isolated from one another and further 
research, including molecular analysis, would very likely bring to light 
diagnosable differences not only between Madeira and Canary birds 
but perhaps also between populations of different Canary Islands. 
Himalayan maximus and Chinese mandarinus would certainly be given 
species status, while the szmillimus group of peninsular India and Sri 
Lanka would probably be treated as three species, as they comprise 
three rather distinct allopatric groups (5 subspecies have been 
recognised, but at least one pair of them intergrade). Central Asian 
intermedius might well be treated as a species, as would the main bulk of 
the west Palaearctic mainland populations within which variation is 
clinal. This clinal variation would not be given formal taxonomic 
recognition (see below). In fact, a major re-examination of the whole 
complex would be necessary before any final decision could be made on 
the division, into a number of phylogenetic species, of what is now 
treated as polytypic Turdus merula. 

The Coal Tit Parus ater would present a similar, perhaps even worse, 
problem. It has a vast continental range in Eurasia in which variation is 
clinal, and a number of isolated peripheral populations, some of which 
are very distinct. At the extreme west of the range, P. a. hibernicus from 
Ireland is diagnosable over most of that island by a yellowish wash over 
its plumage, but some individuals in eastern Ireland are indistinguish- 
able from the British population, britannicus, and a small proportion of 
birds in western Britain show a slight development of the yellow 
colouring that is marked in Irish birds. As British and Irish birds are 
not all diagnosably distinct from one another they would have to be 
treated as conspecific. Furthermore, the Coal Tits from the Iberian 
peninsula are very similar to, and in fact doubtfully distinguishable 
from, British Coal Tits; to the east they intergrade with nominate ater 
which extends over most of Europe and Siberia. At the eastern end of 
the range, there is a population in northern China very distinct from 
nominate ater, with a conspicuous crest and markedly different 
plumage colours (pekinensis). It is not isolated from nominate ater, but 


D. W. Snow 115 Bull. B.O.C. 1997 117(2) 


intergrades with it in Manchuria, Korea and the Amur region. Under 
the PSC all these populations, from Ireland and Iberia east to China, 
would have to be treated as conspecific, and no formal recognition 
would be given to their many and striking differences. 

Attention would then have to be given to the isolated southern 
populations. They comprise what are usually recognised as 14 
subspecies, ranging from northwest Africa east through central Asia to 
southern China and Taiwan. Some are well isolated and distinct, and 
would certainly be treated as phylogenetic species, e.g. cypriotes from 
Cyprus and ptilosus from ‘Taiwan. Others would be problematical. For 
instance, the two northwest African forms, atlas and ledouci, are very 
distinct from all others. They differ conspicuously, but superficially, 
from each other, /edouci from mesic woodlands in Tunisia and eastern 
Algeria having a bright yellow wash over its plumage that is totally 
lacking from the very grey-plumaged, and also slightly larger, atlas 
from the Moroccan Atlas Mountains, which lives mainly in drier 
woodlands and at higher altitudes than /edouci. ‘These two forms have 
stouter bills than their European relatives and differ from them in voice 
(Cramp & Perrins 1993). Under the PSC they would probably, at least 
until a short time ago, be treated as two species. There are, however, 
complications. It has recently been found that some populations 
apparently isolated in patches of rather dry montane woodland in 
Algeria, between atlas and ledouci, are intermediate in plumage but 
closer to atlas in size. They may constitute a third distinct form (Cramp 
& Perrins 1993), but have not been well studied. The Coal Tits from 
the Crimea, where they are well isolated from other populations, are 
usually considered to represent a distinct subspecies, but it 1s not 
well-marked. Further east, from the Caucasus down into the mountains 
of Iran, there is a trend towards brown-backed and _ thick-billed 
populations, apparently adapted to dry oak woodlands. Some at least of 
them are probably well isolated from others by intervening desert or 
semi-desert; but they have not been studied in detail and the three 
subspecies recognised may well need revision. 

It is clear that very considerable research would be needed by anyone 
undertaking to reclassify the Coal Tits according to PSC principles. 
One can only guess at how many species would result; probably at least 
15, and the number might well be altered by later research. 

‘These examples were chosen because I happened to be familiar with 
them, and they may for that reason be suspected of being biased so as to 
exaggerate the problems of using the PSC. To check on this, I made an 
analysis of the 94 European breeding species in the two volumes (5 and 
7) of Birds of the Western Palearctic containing the Blackbird and Coal 
Tit, using the detailed information given in the sections on 
geographical variation, nearly all by C. S. Roselaar. I tried to see how 
many phylogenetic species they would constitute and how many 
problems, and of what kind, would be encountered in attempting such 
a classification. To anticipate what is dealt with in a little more detail 
below, it must be noted again that the PSC gives no taxonomic 
recognition to geographical variation within continuous populations, 
however well marked it may be; all distinct and isolated populations 


D. W. Snow 116 Bull. B.O.C.4997 liv @) 


are, as we have seen, given species status. The analysis gave the 
following results, summarised here in broad categories because hardly 
any two cases are exactly alike. 


No. of spp. 
(BSC) Nature of geographical variation Presumed PSC treatment 
21 None or very slight (monotypic Same number recognised 
species) 
18 Slight; mainly or entirely clinal Ditto; slight geog. variation ignored 
4 Moderate; mainly or entirely clinal Ditto; moderate geog. variation 
ignored 
5 Marked; mainly or entirely clinal Ditto; marked geog. variation 
(e.g. Parus montanus) ignored 
13 Slight, mainly clinal, but with more  Disjunct populations recognised as 
distinct disjunct populations (e.g. species; each BSC sp. probably 


Motacilla cinerea, Evithacus rubecula) becoming 2—4 PSC spp. (further 
study needed) 

8 Moderate or marked, with some Disjunct populations recognised as 
disjunct populations (e.g. Monticola species; each BSC sp. probably 
solitarius, Turdus merula, Parus ater) becoming 2—c. 15 PSC spp. (further 

study needed) 
12 Marked; in distinct subspecies Impossible to predict, would need 
groups, some clinal variation within detailed study 
groups or areas of secondary 
contact/intergradation (e.g. 
Cercotrichas galactotes, Saxicola 
torquata, Parus caeruleus, Lanius 
excubitor) 


is Complex and (except in 2 cases) Ditto 
marked; mainly continental ranges, 
probably with complex evolutionary 
histories (e.g. Galerida cristata, 
Motacilla flava, Sitta europaea, 
Remiz pendulinus) 


The above figures indicate that about half of the species recognised 
under the BSC would need further study before any attempt could be 
made at a PSC classification; many would have to be the subject of 
major revisions. Such research would certainly lead to improved 
understanding of the assemblages of forms involved; it seems unlikely 
that it would lead to stability in species-level nomenclature in the 
coming decades. 

In making this analysis, the strong impression was gained that the 
great majority of polytypic species recognised under the BSC, although 
not based on cladistic analysis, would stand up to such an analysis and 
be shown to be monophyletic. In a few cases, there is doubt about 
whether the species recognised should be split, or merged with another 
species, but not about their monophyly. Before the polytypic species is 
rejected by proponents of the PSC, I suggest that it would be desirable 
to subject a representative selection of widespread polytypic species (as 


D. W. Snow Bz Bull. B.O.C.1997 117(2) 


recognised under the BSC) to cladistic analysis, rather than reject the 
concept wholesale because of detected errors and questionable methods 
previously used in avian taxonomy. 


Recognition of subspecies under the PSC 


It has been widely recognised that the subspecies is not an altogether 
satisfactory taxonomic category, mainly because (1) geographical ranges 
of subspecies cannot be clearly delimited in clinally varying popula- 
tions; (2) there is no general agreement (though some suggestions have 
been made) on what degree of difference between populations should 
be accepted as justifying their subspecific separation; (3) it is very often 
impossible to decide whether or not isolated and _ well-marked 
subspecies should be given specific rank. Means have been suggested 
for dealing with these problems—e.g. Huxley’s (1938) notation )for 
clines; the use of special terms such as semi-species or megasubspecies 
for very distinct forms thought to be verging towards specific 
status—but they have not been generally adopted. Lack (1968) and 
later authors have concluded that the objections are so serious that the 
subspecific terminology at present in use has outlived its usefulness and 
should be replaced by some more appropriate method of categorising 
variation below the species level. 

Under the PSC, the subspecies at present has a shadowy existence 
in a kind of limbo. In his general, theoretical discussion of the 
phylogenetic species, Cracraft (1987) mentions subspecies only in a 
footnote, which is devoted mainly to the problems discussed above and 
implies that the naming of subspecies would have no place in the PSC. 
In his re-classification of the birds-of-paradise (1992), he is more 
explicit, but still with some ambivalence: “‘Adoption of the phylo- 
genetic species concept solves a long-standing source of contention 
within systematics, namely the taxonomic status of subspecies. Because 
phylogenetic species are basal (smallest recognizable) differentiated 
taxonomic units, subspecies could only be applied as arbitrary 
descriptors of within-species variation. Within that context, therefore, 
they serve little useful purpose.” In his Cape Verde Islands check-list 
Hazevoet (1995), after summarising the subspecies problem, concludes 
that “‘trinomials can perhaps still serve a minor role within a continuum 
showing clinal or otherwise geographical variation’’, but whenever he 
uses it he places the word subspecies in inverted commas, implying 
non-recognition of it as a valid taxonomy category. 

Both authors therefore recognise the existence of geographical 
variation within the phylogenetic species, but decide that it does not 
need formal taxonomic recognition. Under the BSC, some kind of 
formal recognition of within-species variation is clearly essential, and 
the subspecies, for all its shortcomings, is the only method that has 
found widespread, though not uncritical, acceptance. If the PSC were 
brought into general use there would surely be a need, in formal listings 
of bird species, to draw attention in some way to clinal variation, in 
which differences between ends of a cline may be as marked as, or in 
some cases considerably greater than, differences between taxa that 


D. W. Snow 118 Bull. B.O.C. 1997 117(2) 


would rank as species. Would it be sensible, for example, to give no 
formal recognition to the striking difference between the small, dark, 
brownish Willow Tits Parus montanus of lowland western Europe and 
the large, very pale, greyish populations of eastern Siberia, because 
they are connected by a range of intermediate forms? 


Use of the PSC in check-lists 


Systematic listing of the avifauna of the Cape Verde Islands (Hazevoet 
1995), with comparatively few breeding species; is not very much 
affected by the choice of species concept. Hazevoet recognises 14 
‘“‘diagnosably distinct endemic taxa’’, which he treats as phylogenetic 
species. Four of them (Apus alexandri, Alauda razae, Acrocephalus 
brevipenms, Passer tagoensis) are distinct enough. to be treated as 
endemic species in Birds of the Western Palearctic, the latest standard 
work covering the Cape Verdes. Three of the remaining 10 are oceanic 
birds (Pterodroma, Calonectris, Puffinus spp.), which present their own 
problems of classification; see, ‘for example, Shirihai et al. (1995) on the 
Puffinus assimilis/Iherminieri ‘complex. The remaining 7 land-birds 
include such forms as Buteo (6.) bannermam, Tyto (a.) detorta and Falco 
(p.) madens, which under the BSC are currently treated as well-marked 
subspecies. 

Hazevoet lists 8 “‘taxa of which the Cape Verde population has been 
treated as a ‘subspecies’, which is, however, not diagnostically 
distinct’. Some of these are certainly very poor subspecies; he 
synonymises them with their respective continental species. But among 
them is the Grey-headed Kingfisher Halcyon leucocephala, which 
differs to a certain extent in coloration and measurements (especially 
bill-length in which there is little overlap) from its Afrotropical 
relatives. It is resident, having lost the migratory habit of the mainland 
populations. Using traditional methods of diagnosis based on 
measurements and plumage, and confining comparisons to adult 
specimens in fresh plumage, it seems quite likely that it would prove to 
be diagnosably distinct. Molecular analysis might be expected to 
support the distinction. In passing it may be suggested that it was by 
good fortune that the other 7 Cape Verde land-bird subspecies all 
proved to be poor ones; applying the PSC to some other island 
avifaunas might have been trickier. 

A case like this suggests that proponents of the PSC, when listing 
local avifaunas, will be confronted with problems similar to those that 
face the conventional BSC taxonomist; but for the former the judgment 
is about the existence or otherwise of a species, and a mistake is more 
serious. 


Application of the PSC to an inadequately known family 


Cracraft’s (1988) re-classification of the birds-of-paradise highlights 
some of the difficulties of applying the PSC to a family in which ranges 
and geographical variation are incompletely known. His analysis results 
in the recognition of 90 species, double the number recognised under 


D. W. Snow 119 Bull. B.O.C. 1997 117(2) 
the BSC (40, Mayr 1962; 42, Gilliard 1969; 45, Sibley & Monroe 1990). 


The increase in number of species comes mainly from the elevation to 
species rank of a large number of BSC subspecies. 

Such a procedure is unjustified in the present state of knowledge. 
The collection and taxonomic study of birds in New Guinea to date 
involves a tiny fraction of the area of the island. In many cases nothing 
is known of the possible presence, or variation, of populations between 
the places where forms usually treated as subspecies have been 
collected. There may well be continuous populations, with intergrada- 
tion; in such cases two or more phylogenetic species would have to be 
merged. Conversely, as Cracraft notes, further collecting may reveal 
new forms, which would be given species status under the PSC if their 
allopatric status vis-a-vis closely related forms could not on available 
evidence be disproved. In all such cases the mistakes that may result 
from inadequate knowledge are more serious for the PSC than for the 
BSC. For instance, Cracraft recognises three species of Paradigalla. A 
recent thorough analysis of this little known genus (Frith & Frith 1997) 
has shown that one of the three, a controversial BSC subspecies based 
on few specimens, is in fact indistinguishable from one of the two other 
(clearly diagnosably distinct) forms. The three PSC species should be 
reduced to two; under the BSC, the only change necessary is to reject 
one doubtful subspecies. 

Cracraft’s treatment seems in some cases inconsistent with PSC 
principles, probably because subjective judgment still seems necessary 
in spite of professions of rigorous analysis. Thus the two populations of 
Lycocorax pyrrhopterus, on Batjan and Halmahera, seem from his 
account to be diagnosably distinct; but he does not treat them as two 
phylogenetic species. The reason (““These two populations are united 
here until further information is available as to the extent of their 
behavioural and genetic differentiation’) surely applies equally to 
forms that he does treat as separate phylogenetic species. 

One result of elevating to species rank very similar subspecies of one 
species within a genus, such as the four forms of Paradisaea minor, is 
that the resulting genus consists of some very distinct and some 
all-but-identical species. There is then an obvious need to recognise a 
category intermediate between the phylogenetic species and the genus, 
as Cracraft does by placing the four above-mentioned forms in a 
‘species group’. This is likely to be a common problem for the PSC. If 
species groups of this sort are formally recognised, the arrangement 
that will result is likely to parallel, and in many cases become identical 
with, the BSC arrangement of polytypic species and their component 
subspecies; the PSC would simply have elevated each category to a 
higher rank. And all this would be due to what seems to me to be a 
dogma rather than a reasoned decision, that species must be the smallest 
diagnosable clusters or end-products of the evolutionary tree. 


Conclusion 


The arguments set out above lead, I suggest, to the conclusion that the 
biological species concept, while not perfect, is still useful and 


D. W. Snow 120 Bull." B.O.C A997 Fae) 


meaningful; and that adoption of the phylogenetic species concept is 
likely to be beset by problems that are equally or more intractable; at 
the very least, they will require years of research in order to establish 
which taxa, now treated as subspecies, have uniquely diagnosable 
characters. Even if this could be done, there would be the risk that 
future research and analysis, using more sophisticated methods, might 
necessitate constant changes. Examples of this are in fact already 
apparent in recent publications. Thus, in their analysis of genetic and 
phenotypic differences between related bird populations on either side 
of the Bering Strait, Zink et al. (1995) find that, on the basis of their 
DNA, the populations of Anthus rubescens would be considered 
different phylogenetic species, but no morphological differences are 
detectable. For some reason they do not do so on the basis of DNA 
alone (perhaps sensing that this would open the floodgates’), but write: 
“If morphological or other genetic differences between these 
populations are found, we think that they should be treated as separate 
species’. How many other such cases are there which may come to 
light; and. how many that will not come to lhght? From a purely 
practical point of view, the adoption of the PSC would mark the end of 
the relative taxonomic stability that has been achieved by the 
acceptance of the biological species concept. From a theoretical point of 
view, its adoption would not, in my opinion, signal any advance in our 
understanding of evolutionary processes at the species level. 


Summary 


The biological species concept (BSC), which has been generally accepted as the basis of 
avian taxonomy for over 50 years, has recently been challenged as being fundamentally 
flawed, and the proposal has been made that it should be replaced by the phylogenetic 
species concept (PSC). Under the PSC, polytypic species are not recognised, the species 
being defined as the smallest diagnosably distinct cluster or population of individuals that 
normally interbreed only among themselves. Reclassification of birds on PSC principles 
would at least double the number of species, but this is not a reason to reject it. 

It is argued here that the concept of the polytypic species is based on recognition of a 
real and widespread phenomenon, and is not invalidated by the old-fashioned and 
inadequate methods that have often been used in its study. The problems encountered 
are just those that would be expected. It is also argued that the definition of what 
constitutes a species under the PSC is arbitrary and includes an element of dogma, and 
has no more inherent validity than definitions based on the BSC. 

Wholesale application of the PSC to birds would meet with serious problems. There 
are many polytypic species (as recognised by the BSC) in which much further, and 
difficult, research would be needed if they were to be reclassified according to PSC 
principles, decisions about what is or is not a diagnosably distinct population being 
impossible on present knowledge. Even if this were done, there would always be the 
likelihood that further research would entail further changes. All such changes under the 
PSC would involve changes in the number of species recognised, and so are more serious 
than changes under the BSC, which would usually affect the recognition of subspecies; a 
category well known to be subject to change. The relative stability in species-level 
taxonomy achieved under the BSC would be lost, certainly for decades. To judge from 
publications to date in which the PSC is adopted, there would still be a need for the sort 
of subjective judgments that have been condemned as a fundamental weakness of the 
BSC. It is concluded that there is at present no case for replacing the BSC by the PSC. 


Acknowledgements 


For help in the preparation of this paper, or criticism of it in draft, I am indebted to 
C. B. Frith, A. G. Knox and J. V. Remsen. 


D. W. Snow 121 Bull. B.O.C. 1997 117(2) 


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Cracraft, J. 1987. Species concepts and the ontogeny of evolution. Biology and Philosophy 
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Address: D. W. Snow, The Old Forge, Wingrave, Aylesbury, Bucks HP22 4PD, U.K. 
© British Ornithologists’ Club 1997 | 


N. F. Collar 122 Bull. B.O.C. 1997 117(2) 
‘Taxonomy and conservation: chicken and egg 
by N. F. Collar 


Taxonomy: a dying discipline in a dying world 


Taxonomy precedes conservation. This is as basic as to say that 
language precedes education. The one is a precondition of the other. 
Without the formal structure of names and an agreed system of usage, 
there can be no understanding of what exists to be conserved. So, if 
taxonomy is simply the time-honoured chicken and conservation her 
newest-laid egg, any implied circularity in their relationship—the point 
about chickens and eggs being that you cannot say which came first— 
might appear whimsical. However, I believe there is an important sense 
in which that circularity—conservation engendering taxonomy no less 
than vice versa—is now a real phenomenon. 

This new emphasis is a side-effect of the widely perceived “‘global 
biodiversity crisis’, in which catastrophic depletion of the natural 
resources of our planet by entirely unsustainable and barely controlled 
patterns of human exploitation—‘‘development”’ is an absurd word for 
it—is engulfing an unprecedented proportion of the world’s animal and 
plant species in a man-made extinction spasm (Wilson 1992, May et al. 
1995). Many (meaning millions) of these species are not yet known to 
science; many (meaning hundreds of thousands) are in the half-way 
house of knowledge between original description and a _ basic 
understanding of their habitats, biology and distributions: up to 40% of 
the world’s 400,000 described species of beetle, for example, are still 
known only from their type-localities (Mawdsley & Stork 1995). The 
utter inadequacy of our knowledge (although of course it has always 
been the case that the exploration of the planet has only ever been a 
subset of its exploitation) is a crippling disadvantage. 

Consequently there is now enormous pressure on the discipline of 
taxonomy, whose services have never more urgently been needed; 
yet in one recent calculation (Soulé 1990), made by multiplying 
the number of tropic-specialised systematists on earth (1500) by the 
number of new species each describes per year (5) and dividing the 
result into the anticipated number of species awaiting description 
(30,000,000), it transpires that on present effort we will need to wait 
until the year 6000 before we have names for all the planet’s species. 
Worse, this “‘present effort’? shows no sign of maintaining itself for the 
next 40 years, let alone the next 4000: 23% of teachers of systematic 
biology in the U.K. were 35 and under in 1980, falling to 8% in 1990 
(Gaston & May 1992). Indeed in the 20-year period after 1975 the 
Sub-department of Ornithology at the British Museum (Natural 
History) lost over half its staff, and more significantly some 90% of its 
research capacity (R. P. Prys-Jones, pers. comm.). This is the 
institution that houses by far the greatest amount of ornithological 
reference material on earth, yet now employs not a single full-time 
in-house taxonomist and has undertaken no exploration or collecting 


N. ¥. Collar 123 Bull. B.O.C. 1997 117(2) 


work of any coherent description since the Mount Nimba enterprise a 
quarter of a century ago (see Colston & Curry-Lindahl 1986). The 
situation is little different in virtually all the great old natural history 
museums of Europe. 

Ironically, although the relative (and I stress: relative) completeness 
of avian taxonomy has prevented it from adding significantly to modern 
museum angst over function, direction and need, the current trend 
(perhaps a better word is thrust) appears to be one that will only add to 
the time it takes for a yet more complete ornithological world picture to 
be composed. Conservationists, however, may not be willing to wait; 
and I am inclined to think that their alliance with birdwatchers—a very 
strong link having been forged between the two in recent years—will be 
at least as crucial to the advance of avian taxonomy in the next few 
decades as any developments within formal academic circles. 


Global conservation and international birding: parallel evolution 


Birds are indeed anomalous amongst major classes or their approximate 
equivalents in the animal and plant kingdoms for the (relatively) high 
degree to which their taxonomies and distributions are known. Such 
knowledge is not, and can never be expected to be, complete, but it is 
largely sufficient for the purposes of global conservation. This is in part 
a reflection of the relatively low total number of recognised species 
(under 10,000). It is also in part a reflection of the relative ease with 
which birds—mostly diurnal, often brightly coloured and normally 
very vocal—are recorded (and indeed adopted as popular objects of 
interest and study). 

There are several important consequences for conservation. First, it 
is actually still possible for an organisation like BirdLife International 
to consider it a realistic ambition to see all the world’s bird species 
saved, even if not by its own hand. Hence the repeated evaluations of 
threatened species in Red Data Books remain relevant, providing the 
stimulus and context not only for single-species initiatives but also for 
site-oriented programmes dealing with major areas of sympatry 
between such species (Collar 1996a). 

Second, the latter principle—targeting areas of sympatry between the 
rarer species, whether technically regarded as threatened or not—can 
be, and has been, formally developed as the basis of more efficient 
conservation investment. This could only be done in a group where 
taxonomic and distributional knowledge was highly advanced. Thus 
ICBP (1992) mapped and overlaid the distributions of 2609 bird 
species with ranges of less than 50,000km7’, resulting in the 
identification of 221 ‘“‘Endemic Bird Areas’? (EBAs). An important 
underlying theory of EBAs is that they are likely also to be areas to 
which other elements of biodiversity are likewise restricted; consider- 
able evidence has been accumulated to support this assumption 
(Thirgood & Heath 1994, Balmford & Long 1995), which therefore 
means that the targeting of conservation resources on EBAs represents 
a considerably more practical alternative to waiting another 4000 years 
for the appearance of a more sophisticated and accurate data set. 


N. F. Collar 124 Bull. B.O.C. 1997 197) 


Third, it has become standard for BirdLife, in various manifes- 
tations, to promote the investigation of many of these threatened 
species, key sites and EBAs via self-funding expeditions, individual 
adventurers, holidaymakers, and of course the ever-increasing numbers 
of in-country ornithologists and conservationists. Biological explor- 
ation, once the exclusive province of museums and run under the 
direction of taxonomists, has transferred to the popular realm of 
conservation and amateur enthusiasm. Between 1983 and 1994, 
BirdLife published the reports of 43 expeditions to 30 different 
countries which its own staff or publications had had some direct 
influence in developing, and stimulated at least as many more whose 
results have appeared elsewhere. There has of course been a 
concomitant growth in interest in wild birds in most countries in the 
world, and BirdLife itself has worked both to encourage such growth 
into coalitions and clubs and to harness them for conservation 
purposes. : 

On the face of it, then, bird taxonomy in the 1990s is not a major 
issue: the species seem largely to have been worked out, and it simply 
remains for their distributions to be more finely plotted. 
Nevertheless—without prejudicing the claim that birds are sufficiently 
well known for small-ranged species to identify biodiversity sites— 
there are hundreds upon hundreds of taxonomic questions relevant to 
bird conservation that remain to be answered, many of them pressingly 
urgent. he interesting part is that, when these questions are not being 
asked by conservationists, and since in Europe they virtually cannot 
now be asked by taxonomists, they are very commonly coming 
instead—often with answers as well—from birdwatchers. 

The reason for this is, of course, that it is birdwatchers who, 
alongside conservationists, are in the vanguard of the new “‘explorers’’, 
moving round the planet long-distance by plane, short-distance by 
logging road, armed with sophisticated optical and tape-recording 
equipment, high-quality protective clothing and the latest immunis- 
ations, and backed up with technical reference material (textbooks, 
recordings) of a totally new level of authority. What we find is that 
more and more such people are coming up against current boundaries 
of taxonomic clarity and knowledge, and are seeking to resolve issues 
beyond these boundaries on the basis of their own increasing 
experience and expertise. They are becoming—one only has to consider 
the burgeoning memberships of the newly founded Oriental, Neo- 
tropical and African Bird Clubs—a new ornithological establishment. 

For such fieldworkers, unallied to museums, the question of 
collecting does not arise. At best, mostly on expeditions, there will be 
mist-netting, which allows detailed descriptions, measurements and 
photographs to be taken. Apart from this, their notebooks are filling up 
with information on foraging habitat and behaviour, nest-site selection, 
general habits and jizz, all of which increasingly feature as evidence in 
the taxonomic decision-making process. Otherwise there remains 
photography and in particular tape-recording. Indeed, it is probably 
mostly in the matter of voice that this army of avian “‘parataxono- 
mists’’, as Janzen et al. (1993) call them, holds the greatest influence. It 


N. #. Collar 125 Bull. B.O.C. 1997 117(2) 


is now common for birdwatchers confidently to pronounce on the 
presence of species on the basis of vocalisations which to the untutored 
ear remain indistinguishable; the rare few, starting with the late Ted 
Parker, have even picked out new species to science merely on this 
feature. 


Species limits: the key issue 


The area of taxonomy which most preoccupies both international 
birdwatchers and conservationists is that of species limits. Generic, 
genetic and other levels of discrimination are largely (at present) 
uncontroversial. ‘The birdwatcher and the conservationist both use the 
species as their unit of currency, and they both therefore have a vested 
interest in knowing what species there are and where, and therefore 
ultimately what a species is. ‘Their natural diffidence over playing at 
taxonomy, which is widely felt to be a demanding full-time 
specialisation, is being forced out of them by the plain unavailability of 
taxonomists, and perhaps also by the growing realisation that a good 
proportion of current taxonomic arrangements is actually the result of 
past amateur (and evidently not always full-time) endeavour. 

Among the obstacles they face is the quality of evidence needed to 
justify the splitting (or indeed lumping) of species. Particularly when a 
form is known to be at significant risk, and if its splitting is justifiable, 
then the case ought never to rest simply on an unsupported opinion, 
and most people would probably agree with (e.g.) Dowsett & 
Dowsett-Lemaire (1993) and Knox (1994) that any such decisions, 
irrespective of conservation considerations, require publication of the 
evidence. This being so, there is additional pressure on the birdwatcher 
and the conservationist to articulate very clearly any views they wish to 
see taken seriously, particularly when there are still taxonomists, most 
now in the U.S.A., well qualified to judge their efforts. This is a 
particularly exasperating circumstance in cases where the original 
professional evaluations under scrutiny (mostly dating from the era of 
taxonomic synthesis in the first half of this century) were themselves 
entirely unexplained and seemingly arbitrary. 

Ideally, therefore, considerable technical detail is desirable, as for 
example has been adduced for Anjouan Scops-owl Otus capnodes 
(Safford 1993) and Wied’s Tyrant-manakin Neopelma_ aurifrons 
(Whitney et al. 1995). On the other hand, time and resources may not 
easily permit such work, so that the rapid articulation of an opinion, as 
in the case of Visayan Flowerpecker Dicaeum haematostictum (Brooks 
et al. 1992), or its containment in a footnote, as with the Chinchipe 
Spinetail Synallaxis chinchipensis (see Ridgely & Tudor 1994), or even 
indeed an assertion as flimsy and unrefereed as that for Sumatran 
Cochoa Cochoa beccaru (Collar & Andrew 1987), may have to suffice. 
Sibley & Monroe (1990) took the unusual step of consulting over 
species limits with fieldworkers, and accepted many splits merely on 
their say-so. This is understandable, given the high degree of 
competence of many birdwatchers and the increasing use of non- 
morphological identification features as taxonomic characters. Indeed, 


N. ¥. Collar 126 Bull. B.O.C. 1997 117(2) 


the great number of further seemingly merited splits, particularly in the 
Oriental and Pacific regions, threatens to occupy so many years in 
documentation that there is clearly a temptation—all the stronger if the 
form is threatened (though not without the danger of distracting from 
other, more certain priorities)—to behave in the same apparently 
cavalier manner as many lumpers did earlier this century. 

The birdwatcher may have more immediate interest than the 
conservationist in splits that involve common forms. The conservation- 
ist will have a far greater interest than the birdwatcher in splits that 
result in significant changes in conservation status; and it is precisely 
because there are potentially major financial and logistical conse- 
quences that such splits should be properly documented, although this 
is not automatically to admit that less rigour is acceptable where 
conservation is not at stake—indeed, a crucial point is that all such 
decisions should be made as far as possible according to the same basic 
criteria. So it matters relatively little to the conservationist that the two 
subspecies of Red-fan Parrot Deroptyus accipitrinus either side of the 
Amazon might better be treated as two species (Whitney 1996), or 
that White-headed and Plum-crowned Parrots Pionus seniloides and 
P. tumultuosus might better be treated as one (O’ Neill & Parker 1977). 
On the other hand, it matters a great deal whether the rare Andean 
representative luchst of the abundant Monk Parakeet Myiopsitta 
monachus has a claim to species status (Lanning 1991), and that 
Indonesia’s Yellow-crested and Australia’s Sulphur-crested Cockatoos 
Cacatua sulphurata and C. galerita might not merit their current 
specific separation, particularly if a decision to lump them (the 
Indonesian form being threatened) were to be accompanied by one to 
elevate the highly distinctive C. s. citrinocristata, confined to the island 
of Sumba. 

It is in particular regard to the fate of such small-island isolates and 
their continental equivalents that conservationists have to be at their 
most sensitive and disciplined, since (a) small-island isolates are 
intrinsically the most susceptible to extinction (King 1985), and (b) 
consistency in decisions over species status is essential for long-term 
confidence and credibility. Thus on the one hand there is pressure, for 
the sake of enhanced conservation interest, to allow specific status to 
distinctive isolates, but on the other there is the inherent inertia of 
established arrangements where the elevation of one subspecies cannot 
wisely proceed without a revision of the entire species or indeed— 
where patterns of subspeciation are judged to reflect biogeographic 
evolution—of an entire local avifauna. 

Some splits, of course, are relatively simple procedures, involving no 
more than two taxa and with no further ramifications, as.in the case 
of the Visayan and Mindanao Broadbills Eurylaimus samarensis and 
E. steertt (split by Lambert 1996) or the Bornean and Sumatran 
Ground-cuckoos Carpococcyx radiatus and C. viridis (split by Collar & 
Long 1995). The problem is less negotiable when concern focuses on 
forms at geographical extremities, which sometimes (owing perhaps to 
founder effects or unusual environmental pressures at the very edge of 
a species’s tolerance) exhibit the most distinctive differences from 


N. F. Collar 127 Bull. B.O.C. 1997 117(2) 


parental stock, for example the curly-tailed, dull-plumaged Tablas 
(Philippines) race menagei (see Vaurie 1949)—alas, probably already 
extinct—of the widespread Asian Spangled Drongo Dicrurus hotten- 
tottus, or the dull red Sumatran race dedemi of the (elsewhere mainly 
green) Grey-headed Woodpecker Picus canus (see Winkler et al. 1995). 
Most problematic of all are highly distinctive forms located 
geographically within a suite of generally less well-marked subspecies, 
like the yellow and red Visayan representative xanthocephalus of the 
normally red, green, black and white Asian Greater Flameback 
Chrysocolaptes lucidus, or the small green Flores form weberi of the 
multicoloured Rainbow Lorikeet Trichoglossus haematodus: splitting in 
such cases could not easily be done without consideration of the 
consequences for species limits in the rest of the complex. These types 
of problem are challenges for those with an interest in order and 
hierarchy and a great deal of time to spare, which is doubtless why so 
many of the earliest taxonomists were aristocrats. The hard-pressed 
conservationist can barely spare two hours, let alone two weeks, simply 
on the effort to establish the taxonomic eligibility of one or two forms to 
join lists of threatened species. 

There are also cases where the effects of taxonomic decisions are 
internal to the complex under review. The Bearded Tachuri Polystictus 
pectoralis, for example, has been judged a near-threatened species 
(Collar et al. 1992, 1994), an evaluation vindicated when a full 
assessment of the evidence was made (Collar & Wege 1995). However, 
the situation only holds while it is agreed that the bird’s three 
subspecies (one Andean, one lowland north of the Amazon, and one 
lowland south of the Amazon) remain as such, since each subspecies, if 
considered separately, would satisfy the new IUCN criteria for 
threatened status (the Andean form may already be extinct). Collar & 
Wege (1995) used this circumstance to illustrate conservation’s 
continuing dependence on taxonomic research and on museum 
collections of birds. In mirror-similar fashion, the New Zealand Brown 
Teal Anas aucklandica meets the new IUCN criteria as threatened only 
while its three races are not upgraded to species level (though this is a 
step already taken by Marchant & Higgins 1990): once they are, two of 
the forms move into a higher threat category than the “‘parent’’ species, 
while the third (the one that continues to take the name A. aucklandica) 
drops out as threatened altogether (the reasons for this are, of course, a 
function of the new criteria and are discussed in Collar et al. 1994: 
19-20). 

In all these cases, of course, we are dealing with a phenomenon— 
allopatric disjunction—with which the biological species concept, 
governed by the principle of reproductive incompatibility in sympatric 
situations, is, on its own admission, ill-equipped to deal. The best it can 
do in this circumstance is to invoke criteria based on certain measures 
of differentiation in related taxa, whether within individual species or 
outside them (see, e.g., Mayr & Ashlock 1991: 104-105); but frequently 
no such taxa exist. Illustration of the resulting variability of treatments 
lies in a review of birds at risk on Negros, Philippines, by Brooks et al. 
(1992). Three of the most threatened are Writh-billed Hornbill Aceros 


N. F. Collar 128 Bull. B.O.C. 1997 117(2) 


(leucocephalus) waldeni, a split recommended by an earlier authority, 
adopted by Sibley & Monroe (1990) and rejected by Dickinson et al. 
(1991) despite the “‘admittedly considerable’’ differences involved; 
White-throated Jungle-flycatcher Rhinomyias (gularis) albigularis, 
lumped by an earlier authority but split by Sibley & Monroe (1990) and 
Dickinson et al. (1991) “on the basis of its plumage differences, 
dependence on lowland forest and disjunct distribution’’; and Dicaeum 
(australe) haematostictum, lumped by Sibley & Monroe (1990) and 
Dickinson et al. (1991) but split by Brooks et al. (1992) themselves 
on the basis of (considerable) plumage differences, unclear vocal 
distinctions and absence of intermediate subspecies. Here then we have 
one major authority (Dickinson et al. 1991) accepting plumage 
differences, allopatry and even habitat to split a species, but unwilling 
to employ the first two of these criteria alone in two other cases, even 
though the differences that mark A. waldeni and D. haematostictum 
from their closest relatives are arguably far more obvious than those 
that so distinguish R. albigulans. 

This variability of taxonomic treatment is an almost inevitable result 
not only of the predispositions and principles of taxonomists 
themselves—in spite of Mayr’s best endeavours over 60 years—but also 
of the patterns of their activity and interest within taxonomy, whereby 
their attention can be concentrated “vertically on particular 
(bio-)geographic groupings (e.g. national or subregional avifaunas), and 
“horizontally” on particular taxonomic groupings (e.g. families, 
genera). The resulting grid tends to leave some groups of species well 
illuminated (not necessarily elucidated, of course) from both directions 
(the Neotropical ovenbirds spring to mind) while others (for example, 
the Oriental babblers) remain in near darkness. The disparities of 
standard, judgement and focus among taxonomists only really emerge 
when their various works, originating at many different points in space 
and time, are synthesised into larger geographical reviews. Indeed, the 
taxonomies of the less worked regional avifaunas commonly display 
inconsistencies based on excessive reverence for stability on the one 
side and overzealous innovativeness on the other. For the conservation- 
ist at the end of the twentieth century, attempting to deal evenhandedly 
with such unevenness can prove to be a decidedly unrewarding and 
lonely task. 


The challenge of the phylogenetic species concept 


The biological species concept (BSC), because of the inherent 
arbitrariness of its treatment of allopatric forms, has been greatly 
vilified as the source of much of this taxonomic doubt and confusion. A 
phylogenetic species concept (PSC), in which a species is simply 
measured as “‘the smallest diagnosable cluster of individual organisms 
within which there is a parental pattern of ancestry and descent’’, is 
being promoted, with increasing insistence, as the one clinically 
objective system that can resolve the problem of allopatric forms (e.g. 
Cracraft 1983, 1992, McKitrick & Zink 1988, Hazevoet 1994, 1996, 
Zink & McKitrick 1995). It is moreover claimed that “because 


N. F. Collar 129 Bull. B.O.C. 1997 117(2) 


phylogenetic species are irreducible, basal taxa, the PSC provides a 
better tool for assessing biological diversity’? (Hazevoet 1994). Clearly, 
therefore, conservationists need to consider the evidence with care. 

The first thing the PSC does 1s obviate the problem of reproductive 
isolation in diagnosable allopatric forms, by pronouncing them all to be 
species; the next thing it does is synonymise BSC _ trinominals 
representative of clines, on the basis that all clinal populations are 
inherently undiagnosable as separate entities. ‘The upshot is an 
anticipated net increase in the total number of bird species and the 
effective total elimination of subspecies. It is, in essence, as simple (and, 
to many, as attractive) as that: the fudge and blur of the BSC 
disappears, and avian diversity emerges as a sanitised, standardised 
construct that allows equal access to all interested parties. 

Sadly, however, the simplicity is illusory. The abiding difficulty of 
this concept is diagnosis. Hazevoet (1994) asserted that there appears to 
be no theoretical limit on the number of species; but this must in part 
be because under the PSC there appears to be no theoretical limit on 
the triviality of the characters used to define them. McKitrick & Zink 
(1988) readily accepted this: 


scrutiny of morphological, behavioral, or biochemical characters on the “‘microgeo- 
graphic”’ level will reveal the existence of many more phylogenetic species . . . There is 
no theory to suggest that a trait must be of a certain quality or magnitude to provide 
historical information or to describe species. - 


On this basis they would admit species status for a population of 
Trumpeter Swans Cygnus buccinator from a ‘‘well-circumscribed 
geographic region”’ if appropriate levels of sampling showed that their 
consistent possession of ‘“‘one extra hooklet on a barb of the seventh 
primary” could diagnose them as an independent evolutionary unit. 
Since they admit the relevance of biochemical characters here, even 
absolute morphological conformity is no sanctuary from the possibility 
of phylogenetic splitting. Cracraft (1992) likewise comments about two 
forms of Manucodia: 


Mayr... notes that trobriandi is smaller than comrii and indeed most specimens fall 
into two distinct size groups... There are, however, some specimens from each form 
that are essentially the same size. Accordingly, only one species is recognized here, but 
future genetic analysis may reveal that the populations are in fact different 
phylogenetic species. 


Here he confirms that diagnosability of characters is indeed a pursuit 
that can draw the taxonomist well below the level of morphology. The 
fact that these two forms happen to show some differences is only 
incidental: biochemistry—despite deepening scepticism in some 
quarters over the efficacy of its methods and hence the validity of its 
results—may legitimately be used to establish phylogenetic species 
even in cases where there are no qualitative morphological indications. 
Hazevoet (1996) makes a similar remark about Pterodroma feae deserta. 

The implications of all this are that many phylogenetic species run 
the permanent risk of representing not “‘irreducible, basal taxa’’ but, on 
the contrary, merely the point at which the search for further 


N. F. Collar 130 Bull. B.O.C. 1997 117(2) 


diagnosable characters has been abandoned. It appears, therefore, that 
we are dealing with a species concept which, like the one it seeks to 
replace and in contrast to the claims of its proponents, can offer no 
actual finality or stability and which, no less than its rival, has 
arbitrariness and subjectivity built into its fabric. Worse yet, we are 
being required to place our trust in its reliance on ever more specialised 
and inaccessible systems of analysis, and we are presumably also 
expected to allow another few centuries while it works its weary way 
through every allopatric population of bird that man can trace and 
scrutinise: every inshore island population of (for example) wren 
(various species of Tvoglodytes), every resident landbird and breeding 
seabird population of every oceanic island (nearly 2000 “‘significant”’ 
such locations listed in Dahl 1991), every montane isolate even within 
the same mountain range, and so on. 

Running parallel is the complex issue of voice and habitat choice as 
characters. It is increasingly frequent for fieldworkers to suggest 
taxonomic revaluations on the basis of these two elements. In particular 
I suspect that because North America leads the world in the realm of 
taxonomy, and because the Americas are so dominated by suboscine 
passerines with their highly stereotyped (i.e. inherited, not learnt) 
vocalisations, the impression across the world is that voice can be 
trusted as a diagnosable character, even though it is well established 
that for the oscine passerines environment—in terms of both 
conspecific singing and local habitat—is a major determinant (Hunter 
& Krebs 1979, Payne 1983, McGregor & ‘Thompson 1988). Thus if the 
Timberline Sparrow Spizella taverneri is allowed species status from 
the allopatric Brewer’s Sparrow S. brewer (Byers et al. 1995) on the 
basis of habitat and voice only, despite its extreme morphological 
similarity, this clearly has implications for any number of other 
allopatric populations of birds which happen to display differences in 
these two parameters: inevitably, again, one is compelled to ask how 
trivial such differences might need to be, and how they might 
objectively be measured. (Indeed, it is time that ornithologists 
considered in depth the entire question of the scientific measurement of 
vocal difference in relation to taxonomic valuation.) 

Scrutiny of an apparently intended model of practical application of 
the PSC—Cracraft’s (1992) review of the Paradisaeidae—only com- 
pounds these uncertainties. This analysis starts with the affirmation 
that, in principle, phylogenetic species are 100% diagnosable, but it 
transpires at once that he commonly lacks the evidence to make his 
judgements conclusive, and thus they often stand as “‘postulations’’; all 
new species, of course, may be considered hypotheses, but the sheer 
degree of provisionality here rather discredits the notion that “a better 
tool for assessing biological diversity’’ is in play. Notably, Cracraft’s 
account of one of two new species his paper describes, Phonygammus 
diamondi, reveals not only the tentativeness of his divisions but also the 
triviality of the characters he allows in making them. In comparing it to 
P. purpureoviolaceus he finds 


the two forms are distinct in that diamondi has the breast and belly dark metallic blue 
with no or relatively little violet-purple suffusion whereas the breast and belly of 


N. 7. Collar 131 Ball. B.O.C.A9971N/(2) 


purpureoviolaceus is strongly suffused with violet-purple. In addition, the lanceolate 
head feathers of diamondi are bluish green in coloration whereas those of 
purpureoviolaceus are greenish blue often with a violet-purple tinge . . . [However] two 
caveats are required. First, there is considerable variation within the taxonomic entity 
purpureoviolaceus and some individuals tend towards diamondi. One specimen of 
diamondi . .. more nearly matches some purpureoviolaceus. [Second] we lack sufficient 
material to say whether diamondi and purpureoviolaceus are really disjunct or whether 
variation between these areas is clinal. 


These admissions hardly confirm the inherent superiority of scientific 
rigour in the PSC; and one’s scepticism only intensifies by reference to 
the start of the paper, where it transpires that the number of specimens 
of diamondi used in its diagnosis was five. This—on the basis of the 
passage quoted—reduces its diagnosability to no more than 80%, so that 
it clearly fails the test of Cracraft’s own criteria, and cannot 
conscionably qualify even as a phylogenetic species, let alone any other 
kind. 

To query the application is not, of course, to invalidate the 
principles, but I think it emerges here how little the PSC differs from 
the BSC in depending, ultimately, on human interpretation of the 
evidence, which at some point inevitably requires reasoned subjective- 
ness. Moreover, it is, I think, legitimate to speculate on its efficacy if it 
performs so inconclusively in the hands of its longest-standing and 
most insistent proponent in ornithology. The 40-42 birds of paradise 
under the BSC become, in this particular exercise, anywhere between 
80 and 120 under the PSC (Cracraft actually established 90, but 
predicted that, while some of these would prove invalid, a good 
proportion of 30 additional races and even some unnamed populations 
would in due course prove ‘““diagnosably distinct’). Extrapolation from 
these figures to the global avifauna produces 20,000—30,000 species, 
leaving no fewer than 10,000 forms whose taxonomic status would 
remain unresolved. Apart from this problem, it appears that the 
admission of increasing levels of triviality opens up the field for 
allcomers to start making their own assessments; where under the BSC 
such parochialism would be contained by the use of subspecies, in this 
scenario full species can in theory be conjured out of next to nothing by 
next to anybody, which will be enduringly contentious, unstable and 
impractical. 

The absence of information on linking populations, of minor 
significance under BSC evaluation, here emerges as crucially important 
to the “‘ontological status”’ (i.e. real-world existence) of a PSC species; 
so for continental isolates, where intervening terrain may hold 
intermediate populations, PSC evaluation must remain provisional and 
hence seriously undependable in the short to medium term. One may 
even ask whether biological diversity is increased or decreased if such 
intervening populations are found to occur: it could not, it seems to me, 
do anything but increase—more populations must mean more 
diversity—yet under the PSC, by its disallowing the taxonomic 
subdivision of clines (which the BSC often arbitrarily breaks down into 
several vaguely bounded subspecies), it would be most likely, curiously, 
to decrease. Conversely, the PSC is exposed to the extraordinary 
situation in which the man-induced extinction of all intermediate 


N. ¥. Collar 132 Bull. B.O.C. 1997 117(2) 


populations in a cline would permit—indeed, I think probably 
require—the diagnosis of the two polarised populations as phylogenetic 
species (an increase in measured biological diversity based on a 
decrease in actual biological diversity). ‘There may be cases where this 
kind of thing is already happening, since any break in a cline of a highly 
sedentary species will isolate common patterns of ancestry and descent 
within the resulting populations. This seems an entirely inappropriate 
yet inescapable upshot—two species, perhaps even three or four if 
enough breaks are made, where recently there was one—whereas under 
-the BSC no change would be expected or registered. 

Conservation, of course, will be more affected than any other 
discipline by further progress of the PSC. Apart from its fundamental 
long-term inconclusiveness, the fact that it would inevitably be very 
patchy in its adoption over time, particularly while it negotiated 
“ontological status’? within the more complex polytypic species, would 
cause serious inequalities in world lists, with parts determined under 
one species concept and parts under another. Then again, even if the 
switch could somehow be effected overnight, there would remain the 
enormous task of distinguishing the threatened species from the secure, 
and of coping with the sheer volume of the former. While it is true that 
some threatened island forms would gain new identity as PSC species, 
the weight of the total number of threatened phylogenetic species (not 
simply proportional to the number of phylogenetic species: Collar 
1996b) would tend to devalue their individual status and “‘jam’’ existing 
priorities with an extensive array of near-lookalikes, drawing attention 
away from the most distinctive species (which, I think, further 
compromises the PSC claim to be “‘a better tool for assessing biological 
diversity’’). Conservationists would then doubtless find themselves 
under pressure (much of it internally generated) to make some kind of 
higher taxonomic prioritisations, which for the most part would lead 
them back towards biological species limits, and which would 
unquestionably absorb a great deal of their very precious time and 
already wilting stamina. 

There is at least one further operational (although not necessarily 
insurmountable) drawback to the PSC for conservationists. ‘This 
concerns the law. For one thing, adoption of the PSC could create 
chaos, at least in the short term, for trade regulation. A PSC revision of 
a family represented on the CITES appendices by many polytypic 
species would, for example, provide an opportunity for traders to test 
the law by trapping all subspecies (now different species) other than the 
nominate, whose name alone would stand as the form (species) 
protected under the convention. Taxonomic updates in CITES take 
several years, and the window of opportunity would be considerable. 
Certain other conventions like the Bonn and Bern likewise possess no 
provisions for rapid taxonomic updates, so again the scope for 
substantial confusion and destabilisation exists; national legislation 
would be similarly vulnerable. As Geist (1992) observed, once names 
are inscribed in law taxonomists forfeit to judges the power to decide 
on matters relating to the definition of species and the criteria for their 
establishment; some unwelcome misalignments could result. Moreover, 


N. ¥. Collar 133 Bull. B.O.C. 1997 117(2) 


in many countries the emergence of hundreds of new species meriting 
protection—these would not necessarily all be globally threatened 
—might prove highly damaging to the cause of those species already 
stretching the capacity and budgets of national conservation agencies. 


Towards a new partnership in avian taxonomy 


The PSC/BSC debate matters to conservation because it has generally 
been standard practice for the species rather than the subspecies to be 
the unit of concern (as is reflected in much national and international 
legislation). The PSC’s greater championing of island isolates and other 
allopatric forms has considerable appeal but, as I have sought to show 
here and elsewhere (Collar 1996b), because of (a) the imbalances it 
would make during the long haul into global application, (b) the logjam 
of (often near-identical) threatened species it would create, and (c) the 
sheer open-endedness of its operation, the PSC appears to have little to 
offer as a conservation tool. My guess is that, once the search for 
diagnosable characters goes biochemical, the entire concept will 
disintegrate as an intended global standard. 

Nevertheless, understandable dissatisfaction with the way lumping 
can mask the “‘true’’ status of various forms (e.g. Whitney et al. 1995) 
requires a response. It seems to me that there are many cases where 
lumping was justified and some where it was not, and that the prime 
task is to concentrate both field and museum studies as much as 
possible on likely instances of the latter, and not to allow frustration 
with the misapplication of one concept to result in complete 
dependence on another. So while the exploration of unknown or 
recently unvisited areas is a continuing ideal of much modern field 
ornithology, the rechecking of ‘“‘subspecies’’—all well-marked (or 
indeed vocally distinctive) allopatric forms—for the teasing out of false 
polytypic amalgams is a parallel incumbency. Objective criteria by 
which such forms should be identified and rechecked are themselves 
in need of development beyond those in Mayr & Ashlock (1991): 
I would imagine that habitat and elevation, and perhaps even 
biochemistry, would normally only be used as supplementary elements, 
but a way must now be found formally to invest vocalisations with the 
same potential taxonomic relevance (and I repeat there are still 
important caveats here too) that morphology has until now exclusively 
enjoyed. 

A second step is the greater familiarisation of dedicated amateur 
birdwatchers with the principles and practices of taxonomy, which is 
something that both taxonomists and conservationists could encourage 
(perhaps a cheap manual would help demythologise the subject). Such 
people are, after all, the representatives of a greater, more general body 
of nature lovers and wildlife enthusiasts to whom conservation and, to 
some extent at least, modern taxonomy owe their existence. The 
democratisation of taxonomy, through the involvement of the growing 
body of highly skilled field experts, would harness new forces in the 
rejuvenation of this most essential and basic of biological disciplines 
(although I should emphasise, particularly in respect of an earlier 


N. ¥. Collar 134 Bull. B.O.C. 1997 117(2) 


concern about next to anybody being able to conjure up phylogenetic 
species, that I see taxonomic democracy as a privilege and 
responsibility that the entire ornithological community should commit 
itself jealously to guard). 

Third, the challenge can partly be met by considered input into 
national conservation strategies, which can and should involve the 
identification (by national agencies) of globally threatened subspecies 
(Collar 1987). These forms can be allotted independent attention 
(including investigation of taxonomic status) chiefly when they prove 
not to be sympatric with “‘endemic”’ or threatened species in areas such 
as those identified by ICBP (1992) and Wege & Long (1995), i.e. when 
they fail to find sanctuary within the main suite of a country’s 
conservation target areas. Such an approach at least promotes vigilance 
for the more vulnerable among less differentiated forms, without 
compromising either the standard use of the BSC or the global 
priorities that result from such use. 

This agenda gives a new impetus to the chicken-and-egg cycle of 
(expressly avian) taxonomy and conservation. Clearly it requires 
coordination and consistency to be effective, and in displaying these 
things it might make its most valuable contribution to the growth of 
relations between taxonomists and conservationists concerned with 
non-avian life-forms, where neither side can boast a distinguished track 
record in strategic planning to optimise the value and impact of its 
work (Mound & Gaston 1993, Collar 1994). Such is the weakness of 
both taxonomic and conservation knowledge in most of these life-forms 
that conflict of interests is permanently likely, especially if taxonomy 
becomes thought of as a service industry whose costs should be covered 
by conservation or at least by the sources that fund conservation. 
Indeed, this is already happening: cutbacks in the core budget of the 
former British Museum (Natural History) have already partially been 
offset by its eligibility to compete, in humiliating sit-up-and-beg 
fashion under its new downsized identity (““Natural History 
Museum’’), for support from the “‘Darwin Initiative’ (the British 
government’s Earth Summit contribution towards the conservation of 
global biodiversity); while the recent Systematics 2000 initiative (for 
which see, e.g., Cracraft 1995) shows every sign of developing into a 
major competitor for big-time biodiversity funding elsewhere in the 
world. 

What is needed in this situation is redoubled government support for 
the institutions of taxonomy (not just in the U.K. but in every country 
with a museum tradition) coupled with a requirement that, rather than 
just resuming their timeless remit to fill out tiny random areas in the 
global biodiversity patchwork, they select and plan out their research 
priorities and activities in a serious, sustained partnership with 
conservationists, to ensure adequate and representative sampling across 
the planet within a time-frame (the next quarter-century) to be of 
maximum benefit to the preservation of biological diversity. The more 
such initiatives are modelled on the use of manageable, relevant subsets 
like those used by BirdLife in its threatened and endemic species 
analyses, the greater the chances that taxonomy can make a genuine 


N. ¥. Collar 135 Bull. B.O.C. 1997 117(2) 


contribution to the process of preserving the largest possible number of 
the earth’s current complement of animal and plant species. 


Acknowledgements 


I am most grateful to Robert Cheke for the invitation to prepare this paper, to Jurgen 
Haffer and two referees for their comments on the typescript, and to Kevin Gaston, Bob 
May, Robert Prys-Jones and Effie Warr for help in the provision of reading material. 


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Address: N. J. Collar, BirdLife International, Wellbrook Court, Girton Road, Cambridge 
CB3 ONA, U.K. 


© British Ornithologists’ Club 1997 


te wie. 


‘ . j 
iat 28 V4 


7, Queiwiks afte 


Plate 2. Protagonists in the trinomial controversy in Italian ornithology. Top: Tommaso 
Adlard Salvadori (courtesy of the Salvadori Muzzarelli Family). Lower left: Enrico 
Hillyer Giglioli (from: Annali di Agricoltura, no. 268, 1912). Lower right: Ettore 
Arrigoni degli Oddi (courtesy of Donna Fabrizia Ruffo di Calabria). 


C. G. Violant & F. Barbagli 137 Bull. B.O.C. 1997 117(2) 


Salvadori, Giglioli and Arrigoni: aspects of 
the trinomial controversy in Italian 
ornithology 


by Carlo G. Violam & Fausto Barbagli 


After the publication of the 10th edition of Systema Naturae (1758) by 
Linnaeus, the adoption of his zoological nomenclature was favoured 
by several eminent naturalists of the 18th century in Italy, such as 
Giovanni Antonio Scopoli (1723-1788) and the Abbé Giovanni Ignazio 
Molina (1740-1829). The former, as a correspondent of Linnaeus, 
started using the binomial nomenclature in Entomologia Carniolica 
(1763) and, later, in 1769, as far as ornithology is concerned, in Annus 
I. Historico-Naturalis. He was the first author to describe the Little 
Owl Strix noctua, the Barn Owl Strix alba and the White-fronted 
Goose Branta albifrons, amongst other birds (these and other scientific 
names here given in their original form). 

The second naturalist, Giovanni Ignazio Molina, was the librarian of 
the Jesuit College in Santiago de Chile; he arrived in Italy after the 
expulsion of the Jesuits from Chile in 1767 and lived in Bologna for the 
rest of his life, where he was ordained priest. In his famous essay 
““Saggio sulla storia naturale del Chili’’ (1782) he described for the first 
time the Chilean Flamingo Phaenicopterus Chilensis, the Black-necked 
Swan Anas Melancorypha, the Patagonian Kelp Goose Anas Hybrida 
and other new bird species, as well as, among mammals, the Coypu 
Mus Coypus, basing the diagnoses on his recollections. However, unlike 
Scopoli, Molina was not a whole- hearted supporter of Linnaeus. In the 
introduction of his ‘“‘Saggio’’ he admitted: 


“*... [have conformed to the Swedish Naturalist, not because I believe that his system is 
superior to all the others, but because I see that at present it is almost universally 
followed. Although my esteem for his knowledge is great, I cannot help saying that I do 
not like his very clever nomenclature in many essential points. I would have rather 
followed Wallerius or Bomare in Mineralogy, the great Tournefort in Botany, and 
Brisson in Zoology, as they seem to me easier and more appropriate for the common 
comprehension ...”’ [transl.] (Molina 1782). 


By the beginning of the 19th century, the use of binomial 
nomenclature was consolidated thanks to Paolo Savi (1798-1871) and 
Carlo Luciano Bonaparte (1803-1857). At the instigation and 
leadership of Prince Bonaparte, the Quarta Riunione degli Scienziati 
Italiani (Fourth Congress of Italian Scientists) was held in Padua in 
September 1842, and on this occasion the ‘Strickland Code’’ was 
presented to the Italian scientific community of the day, and was duly 
translated into Italian for the following year’s meeting at Lucca. The 
use of the “Strickland Code’’, which had been successfully proposed 
during the Manchester Congress of the British Association for the 
Advancement of Science in 1842, aimed at unifying the rules of 
nomenclature, applying the law of priority and sanctioning any possible 
changes when considered necessary. The regulations fixed the 12th 


C. G. Violani & F. Barbagli 138 Bull. B.O.C91997 IWh7@) 


Edition of Linnaeus’ Systema Naturae (1766) as the starting point for 
nomenclature; the Code also had the task of preventing the kind of 
nomenclatural anarchy promoted by zoologists such as Illiger and 
Swainson (Stresemann 1975). During the following years, exploratory 
voyages and the systematic collecting of zoological material started the 
golden era of descriptive zoology, which saw Tommaso Salvadori 
(1835-1923) and Enrico Hillyer Giglioli (1845-1909) as its most 
distinguished representatives in Italian ornithology. 

At Pisa University both scientists, though in slightly different times, 
had been pupils of Professor Paolo Savi, whose concept of nomen- 
clature is clearly expressed in his “Ornitologia Toscana’’ (1827-1831): 


“*... Names should be considered as indisputable, and they must not be changed for any 
reason at all; the first name which has been bestowed on a species is its true one, and it 
must be kept consistently and carefully; in case that the same species—due to the progress 
of science—needs to be placed in another genus, only then should a change in the generic 
name be permitted; the specific name however shall remain unchanged. . .”’ [transl.]. 


Because of their scientific influence, and the fact that they were in 
regular contact with the British school (Philip Lutley Sclater, Alfred 
Newton and Richard Bowdler Sharpe), Giglioli and Salvadori were 
soon considered the most authoritative ornithologists in Italy. 
Furthermore, Giglioli had studied at the Royal School of Mines in 
London and was in close contact with the cultural entourage 
surrounding Darwin and Huxley at the time of the great debate on the 
origin of species (Barbagli & Violani 1996). The work of Giglioli both 
as author and as museologist was based on the study of adequate series 
of specimens, for a better understanding of zoogeography in the light of 
the evolutionary theories. 

In Germany the ideas of Otto Kleinschmidt, followed in Austria by 
Victor von ‘l’schusi zu Schmidhoffen, and in Great Britain by Ernst 
Hartert, soon began to gain ground; these authors started to describe 
subspecies and used trinomials, whose definition was summarized by 
Hartert in “‘V6gel der Paldarktischen Fauna’”’ (1903): 


“*.. . We describe as subspecies the geographically separated forms of one and the same 
type, which taken together make up a species. Therefore not just a small number of 
differences, but differences combined with geographic separation, permit us to determine 
a form as subspecies, naturally when there is general agreement of the main characters 
AE itaraas le 


The scientific establishment in England dominated by Sclater with 
Newton, Saunders, Salvin and Sharpe, was firmly opposed to such a 
definition and to the introduction of a trinomial nomenclature (see 
Stresemann 1975, Haffer 1992). In Italy too, Giglioli disagreed with 
Hartert, as can be seen from his famous speech delivered in 1908 in 
Florence, during the inauguration of the Central Collection of Italian 
Vertebrates; its translated quotes include: 


“,. At this moment when a wind of analysis at all costs is blowing among the 
connoisseurs of natural sciences, while synthesis is either neglected or people do not know 
how to do it, a great danger for the descriptive and systematic aspect of science rises with 
the institution, let us say so, of the subspecies as new forms are being created with new 


C. G. Violani & F. Barbagli 139 Bull. B.O.C. 1997 117(2) 


names willy-nilly more, I fear, to satisfy puerile vanity, than to establish new facts and 
report new cases. This, in addition to the very minute description of individuals, the 
confusion of individual characteristics with specific characteristics, is fragmenting wrongly 
even the common and more clearly distinguishable species, and multiplying scientific 
names in an incredible way; and, I am very much afraid, will lead us to total chaos... . 
If, as I always have, one accepts Darwin’s grand hypothesis, one must recognize that an 
absolute definition of the species is no longer possible; what naturalist would nowadays 
attempt to give the quantity of the species? Nevertheless, the species exists in nature, as 
no one can seriously deny; but it is variable within certain limits, and is not as was once 
believed, a fixed and determinate entity. It is hence obvious that some species are more 
distinct, others are Jess so; the former are further removed from the congeneric species, 
while the latter, which are less well-defined, remain close—sometimes very close—to one 
or more congeneric species, being, nevertheless always detached; under normal 
circumstances, the former will have moved away from the ancestor spécies sooner than 
the latter. According to some, the latter hence constitute the so called subspecies, while 
others see the subspecies as a lesser entity lying somewhere between the species and the 
individual. 

I would agree with the second definition, but it must be clearly stated that these 
subspecies are not entirely detached from the species they derive from: that is to say, 
some individuals will have the intermediate characteristics both of the parent species and 
of the new form in evolution. I could cite not a few examples of such cases; for obvious 
reasons, to these and these only, and even then with a great deal of circumspection, 
should trinomial nomenclature be applied. In my opinion the danger lies precisely in the 
incorrect definition of the subspecies and in the abuse of trinomial nomenclature to name 
the species mentioned above, which are detached from, but very similar to other 
congeneric species; therefore I use binomial nomenclature for these, since one has to be 
careful not to distort the grand, simple and natural Linnean concept, expressing itself in 
its binomial nomenclature. 

... | would like to point out that, besides the damage due to the useless multiplication 
of terms, the trinomial nomenclature would necessarily imply the concept of descent, and 
I do not think it is easy to deduce, in most cases, which is the parent form and which the 
daughter. Therefore when I speak of subspecies, or geographical races, or incipient species, 
that is, species which are still developing, I mean the forms which have varied in a certain 
sense, due to the effects of the environment and of pressure from their ancestor, e.g. the 
species which they derive from, and they pertain to, for the presence of individuals with 
intermediate characteristics connecting them to that species. Since these subspecies are 
not yet separate from the mother species, they cannot be treated or enumerated 
separately, and it is only to them that trinomial nomenclature should be applied. I hope 
I made myself clear on this important matter. The subject deserves this. Hybrids and 
individual variations are, of course, a different matter, not to be confused with subspecies 
or incipient species, much less with the cases of neogenesis which, in my opinion, can 
give rise to a real and proper species, distinctly detached from its parent species. It must 
be remembered, furthermore, that in zoology as in the other biological sciences, 
nomenclature is a means, not an end. Today, the very same blessed nomenclature coined 
by Linnaeus to facilitate and clarify the work of the naturalist has become the greatest 
obstacle and the greatest source of confusion to scholars. I would add that the continual 
changing of the names long used up until now for common species and genera in favour 
of unknown or forgotten names, with the excuse of rigidly applying the law of priority, as 
well as the consequent repetition of the same name for the generic, the specific and the 
subspecific, has reached an intolerable paroxysm; not to mention that such puerile 
incongruity is revolting to even the most basic common sense. As regards myself I am 
also opposed to the simple repetition of the same name with generic and specific value. 
The enormous damage to the confusion in zoological nomenclature, to which some have 
even tried to make the great Linnaeus himself accomplice by using the Xth edition of 
Systema Naturae rather than the classical XIIth edition amended and corrected by 
himself, rightly used as the source for binomial nomenclature, prompted the discussion of 
a motion in section D (zoology) at the recent meeting in Dublin of the British Association 
for the Advancement of Science, to establish that the names in general use and which 
were used by the great biologists of the last century be unalterable. The proposal was 
therefore backed by the most illustrious British zoologists. (Cf. Nature, 27 August 1908). 
Even in the United States, where much has been committed in the name of trinomial 
nomenclature, last spring saw the beginning of an attempt to ward off the serious danger 


C. G. Violani & F. Barbagli 140 Bull. B.O.C. 1997 117(2) 


caused by the excessive multiplication of forms with specific value based on insufficient 
characteristics. (Cf. American Naturalist, April 1908).”’ [transl.]. 


Elsewhere, in the last volume of his “Inchiesta ornitologica’’, Giglioli 
(1907) published some witty remarks, always on the same theme. Two 
of these, on the Corsican Nuthatch Sztta whiteheadi and on the Jay 
Garrulus glandarius, are particularly interesting in this respect. While 
dealing with S. whitehead: he argued with the bird curator of the 
Rothschild Museum: 


“‘... Hartert considers the Sztta whiteheadi as a mere subspecies of Sitta canadensis, to 
which it is doubtless remarkably similar; but in this case it seems to me that my friend has 
been driven by his fatherly love for subspecies.’’ [transl.]. 


Even for the Jay, Giglioli disagreed with his transalpine colleagues; 
after the comparison of hundreds of specimens obtained from 
everywhere in Italy, he became convinced that it was a very variable 
species and therefore it was impossible to distinguish not only a 
subspecies, but even some “‘local varieties” of it. So he specified: 


““’.. For this reason I believe it simply absurd to have separated, even subspecifically, 
jays collected in Sardinia from those of our mainland, as Kleinschmidt, Tschusi and— 
unfortunately—the able Hartert have done. And so, at a distance of one month, the 
former author christened his great discovery as Garrulus ichnusae, the latter one as 
Garrulus glandarius sardus! 1 feel I am watching a puerile competition on a greasy 
pole! . (transla: 


The thoughts of Salvadori ran along similar lines, but emerged in 
less emphatic tones, mainly in private documents such as his 
correspondence with R. B. Sharpe. In a letter dated 10th March 1904, 
written in a spell of low spirits, he blamed the German authors: 


“«.. I must say that I am rather disgusted with the recent work of many ornithologists. 
When the trinomial system has the consequence of adopting such names as Coccothraustes 
coccothraustes coccothraustes, Oriolus ortiolus oriolus, Pica pica pica, I conclude that the 
system is absurd! No less absurd is Hartert’s system of using the names Carpodacus 
(mas.) rhodochroa (fem.), C. rhodopepla, C. rosea, C. erythrina, C. synoica. Sensible people 
will laugh at us if we continue in this way ...’’ (Violani in: Anon. 1993). 


In the paper written for The Ibis Salvadori (1904) carried on a 
controversy with his friend Hartert concerning the choice of the 10th 
edition of Linnaeus’ Systema Naturae as the starting point for 
zoological nomenclature and its consequences in the case of the Latin 
naming of two common species of European thrushes, Turdus musicus 
and Turdus tliacus. 

The first Italian author of the new generation, 22 years and 32 years 
younger than Giglioli and Salvadori respectively, who did not oppose 
the trinomial nomenclature is Count Ettore Arrigoni degli Oddi 
(1867-1942), owner of an extraordinarily rich private collection of birds 
(Barbagli et al. 1996), and in touch with Tschusi and Hartert as a 
collector. He adopted trinomial nomenclature, when in 1902 he 
described seven subspecies of birds (five from Sardinia, one from 
Corsica, one from mainland Italy), thus publishing avian trinomial 
names for the first time in Italy (Arrigoni degli Oddi 1902a, 1902b). 


C. G. Violani & F. Barbagli 141 Bull. B.O.C. 1997 117(2) 


Although Arrigoni degli Oddi had joined the new school as far as 
trinomial nomenclature was concerned and used it in his “‘Elenco degli 
uccelli italiani’ (1913), in the same book he made it clear that: 


*«... As regards subspecies, I did not omit to mention them under each single species, mostly 
in order to keep my work updated; as for myself in most cases I am not even able to distinguish 
them and I consider an analysis carried to the extremes to be a very serious damage to science. 
With the greatest thoughtlessness new names are given to new forms almost always based on 
individual, not specific, characters; in this way, such a confusion of technical names and of 
forms is created, that sometimes one is at a complete loss . . .”’ [transl.]. 


The same concept was taken up again in greater detail by Arrigoni degli 
Oddi in “Ornitologia italiana’? (1929), when the author explained his 
objection to the contemporary significance and usage of the subspecies: 


“... The name “subspecies” was given, adopting trinomials, to those small variations 
which depend on habits, life environment, locality or other factors, emanating from a 
close origin, and, if not in all, consistent in their complex and to which they are connected 
thanks to intermediate individuals. Today, however, the subspecific distinctions multiply 
continuously, and in such way that they give birth to great confusions, and make scientific 
researches less serious. Different names were also proposed in order to indicate the 
various modifications undergone by the type species in the different countries, and 
excessively fanciful or unimportant features were fixed. ‘““This according to myself [M. 
Paulucci 1879, p. 79] cannot be undertaken seriously, as these modifications pass 
imperceptibly from one to the other, and through innumerable tiny changes unrolls and 
develops a chain formed by a quantity of small links all connected together.’’ Many of 
these modern distinctions were established upon a single specimen, comparing it with 
some similar ones from a very distant locality, without paying attention to the fact that in 
the intermediate countries the various links could be found, and without considering that 
it is with large series that species must be established. . .”’ [transl.]. 


As far as trinomial nomenclature was concerned, the controversy in 
Italy was a matter more of form than of content; indeed, during his 
studies on the Moluccan and Papuan avifaunas Salvadori did describe a 
Semioptera wallace: var. Halmaherae, as well as a Paradisaea apoda var. 
novae guineae, signifying his intention to point out morphological 
differences when perceptible and consistent (Salvadori & D’Albertis 
1879, Salvadori 1882). 

In reality, Salvadori rather often employed the term ‘“‘variety’’ in 
order to indicate individual variations, as for instance Eos riciniata 
(Bechst.) var. cauda viridi (Salvadori 1881). In the above cases of birds 
of paradise, however, as is shown by the geographical names adopted 
and by the considerable number of syntypes examined (40 specimens of 
var. novae guineae and 50 of var. Halmaherae), he intended to designate 
two entire morphologically consistent local populations; thus he used 
the term ‘“‘variety’ in the same way as the new school of his time 
was using the word “‘subspecies’’. The step between ‘“‘variety’’? and 
“‘subspecies’’ was short, as is apparent from the ruling of the I.C.Z.N. 
(1985) to consider the term ‘“‘variety’’ as of subspecific rank, if 
published before 1961, unless the content of the work reveals that 
infrasubspecific rank is meant [Art. 45(g)]. 

The reluctance of Giglioli to use “‘subspecies’’ derived from the 
impossibility of arranging trinomials according to phylogenetic descent; 
consequently, he indicated as species different forms even ‘“‘very little 


C. G. Violani & F. Barbagli 142 Bull. B.OvCyl997 ATIC) 


detached’? from each other, on condition that all the individuals 
belonging to these forms could be separated on the basis of 
morphological characters. His belief is well expressed in a passage on 
the Black-eared Wheatear Saxicola melanoleuca (now Ocenanthe 
hispanica melanoleuca Guldst.). 


““... The difference between these two forms [,Saxicola melanoleuca and S. occidentalis| is 
not great, but it is constant, therefore they should be kept as distinct. According to 
modern criteria they must be considered as subspecies or incipient species, as it is better 
expressed; but it is not possible to tell which of the two is the more ancient, or from what 
ancestor the other has derived. Such a fact illustrates very clearly the fallacy of burdening 
the already overloaded ornithological terminology with a trinomial nomenclature. Who 
could prove to me in this case (and almost all the so-called subspecies are in such a 
condition) that it is correct to write: Saxicola occidentalis melanoleuca or Saxicola 
melanoleuca occidentalis, in order to distinguish the two forms; or, on the contrary, using 
the incongruous and monstrous repetition of the specific name in order to designate the 
mother form, to write: Saxicola melanoleuca melanoleuca or MSaxicola occidentalis 
occidentalis? For obvious reasons of order and logic, I prefer to maintain the admirable 
and very simple binomial nomenclature of the most celebrated Linnaeus; affinities, as 
well as divergencies between two forms (let us call them “‘species’’), are variable 
quantities and easily measured very differently by the various scholars, so that we cannot 
always have a sharp and precise definition to distinguish species from subspecies; still less 
easy is it to establish which is, between two related forms, the (mother) species, or the 
(daughter) subspecies; since such should be their true and phylogenetic relationships .. .”’ 
[transl.] (Giglioli 1894). 


After the death of Giglioli and Salvadori, the use of trinomial nomen- 
clature in ornithology was accepted in Italy without further controversy; 
Arrigoni degli Oddi used trinomials in ‘his textbook ‘“Ornitologia 
italiana’”’ (1929), though remaining sceptical, as we have seen, to the so 
called ‘‘manufacturing of subspecies’’ by his German colleagues. 

As a curiosity it may be recalled that, during the 1950s and 1960s, 
Antonino Trischitta (1892-1966), an eccentric Sicilian naturalist, 
proposed a bizarre nomenclatural system, which he called “‘Nomen- 
clatura Buiologica ternaria plurinominale’ (Ternary plurinominal 
biological nomenclature). This system (Trischitta 1950, 1967) was 
intended to reveal the exact position of the living being, which the 
name referred to, within the systematic frame. The three terms 
indicate: the first, the kingdom, the phylum, the class and order which 
the organism belongs to; the second, the family and the genus included 
in the same family; the third, the species and, possibly, the subspecies. 
For instance, applying ‘Trischitta’s system the scientific name of 
Bonelli’s Eagle would be: 


Zochoaviaccipitrides accipitrihieraaetus fasciatus 


resulting from: 


Regnum: Animale (Zoo) Zor+ 

Phylum: Chordata Zocho+ 

Classis: Aves Zochoavit 

Ordo: Accipitriformes Zochoaviaccipitrides 
Familia: Accipitridae accipitrit + 

Genus: Hieraaetus accipitriihieraaetus 


Species: fasciatus fasciatus 


C. G. Violani & F. Barbagli 143 Bull. B.O.C. 1997 117(2) 


This method had no followers, but nevertheless it stimulated a short- 
lived interest and its mechanism was published in the pages of the 
authoritative “‘Bollettino di Zoologia” (‘Trischitta 1952) edited by the 
Unione Zoologica Italiana. 


Summary 


After the publication of Linnaeus’ Systema Naturae (1758) binomial nomenclature was 
also introduced in Italy and adopted by several famous naturalists of the 18th century, 
such as Giovanni Antonio Scopoli and the Abbé Giovanni Ignazio Molina. By the 
beginning of the 19th century its use was consolidated thanks to the work of Paolo Savi 
and Carlo Luciano Bonaparte. Under the leadership of the latter scientist the Quarta 
Riunione degli Scienziati Italiani was held in Padua in September 1842 and, on this 
occasion, the Strickland Code, subsequently translated into Italian, was recommended 
since it fixed the 12th edition of Systema Naturae as starting point for scientific 
nomenclature. 

Subsequently, voyages of exploration and the systematic collecting of zoological 
material started the golden era of descriptive zoology, which had Tommaso Salvadori 
(1835-1923) and Enrico Hillyer Giglioli (1845-1909) as its most distinguished 
representatives in Italian ornithology. Between the 19th and the 20th century the 
German-speaking ornithologists Ernst Hartert, Otto Kleinschmidt and Victor von 
Tschusi introduced the subspecies concept and began to use trinomial nomenclature 
consistently in their writings. In Italy this new school found an eminent supporter in 
Count Ettore Arrigoni degli Oddi (1867-1942), who in 1902 described his first new 
subspecies of Italian birds using trinomials, in conflict with the old school of Salvadori 
and Giglioli. 

At about the same time in England a similar situation occurred, between the younger 
Hartert and the older ornithologists such as Richard Bowdler Sharpe, Philip Lutley 
Sclater and Alfred Newton. The controversy which ensued lasted for about twenty years 
and is still remembered thanks to private correspondence and articles in scien- 
tific periodicals by the authors involved; though sometimes extremely polemic, their tone 
was frequently enlivened by humour as is shown, for instance, in Giglioli’s writings. 


Acknowledgements 


The authors are greatly indebted to the Linnean Society of London and to the British 
Ornithologists’ Club for their kind invitation and hospitality and for their constant 
encouragement: in particular, they would like to thank Miss Gina Douglas, Mrs 
Amberley Moore, Dr John Marsden, Dr Robert Cheke, Cdr Michael B. Casement and 
Mr David Pescod. Fausto Barbagli is especially grateful for the financial support during 
his visit to Great Britain. Jurgen Haffer has kindly provided relevant literature on the 
controversy in Germany and Miss Claire Archibald, University of Pavia, helped with the 
translation of some parts of Giglioli’s text. 


References: 

Anon. 1993. Report of the Committee for the Club’s Centenary year 1992. Bull. Brit. 
Orn. Cl. 113: 1-9. 

Arrigoni degli Oddi, E. 1902a. Atlante Ornitologico, Uccelli Europei con notizie d’indole 
generale e particolare. Hoepli, Milano. 

Arrigoni degli Oddi, E. 1902b. Note sopra alcune nuove sottospecie osservate negli 
Uccelli di Sardegna. Avicula 7: 102-105. 

Arrigoni degli Oddi, E. 1913. Elenco degli Uccelli Italiani, per conoscere a prima vista lo 
stato esatto di ogni specie, riveduto al 31 dicembre 1912. Bollettino ufficiale del 
Ministero di Agricoltura, Industria e Commercio 12: 1-122. 

Arrigoni degli Oddi, E. 1929. Ornitologia Italiana. Hoepli, Milano. 

Barbagli, F. & Violani, C. 1996. L’ “Inchiesta Ornitologica’’ di Enrico Hillyer Giglioli. 
Riv. ital. Orn . 65: 136-146. 

Barbagli, F., Carlini, R. & Violani, C. 1996. Ettore Arrigoni degli Oddi: cenni biografici. 
Ric. Biol. Selv. 97: 5-26, 307-310. 


C. G. Violani & F. Barbagli 144 Bull. B.O.C. 1997 117(2) 


Giglioli, E. H. 1894. Monachella a collo nero. Saxicola melanoleuca, Guldst. In: Giglioli, 
E. H. & Manzella, A. 1879-1907. Iconografia della Avifauna Italica, ovvero tavole 
illustranti le specie di uccelli che trovansi in Italia con brevi descrizioni e note. Tip. 
Pellas, Firenze. 

Giglioli, E. H. 1907. Secondo resoconto dei Risultati della Inchiesta Ornitologica in Italia. 
Stab. Tip. S. Giuseppe, Firenze. 

Giglioli, E. H. 1908. La collezione centrale degli animali vertebrati italiani nel Regio 
Museo zoologico di Firenze. Atti Soc. ital. Progr. Sc. 2: 1-19. 

Haffer, J. 1992. The history of species concepts and species limits in ornithology. Bull. 
Brit. Orn. Cl. Centenary Suppl. 112A: 107-158. 

Hartert, E. 1903. Die Vogel der Paldarktischen Fauna. Band I. Verlag von R. Friedlander, 
Berlin. 

1.C.Z.N. (International Commission on Zoological Nomenclature) 1985. International 
Code of Zoological Nomenclature. International Trust for Zoological Nomenclature, 
London. 

Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum Classes, Genera, 
Species cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima, 
Reformata. Holmiae, Laurentii Salvii. 

Linnaeus, C. 1766. Systema Naturae per Regna Tria Naturae, secundum Classes, Genera, 
Species cum Characteribus, Differentis, Synonymis, Locis. Tomus I. Editio Duodecima, 
Reformata. Holmiae, Laurentii Salvii. 

Molina, G. I. 1782. Saggio sulla storia naturale del Chilt. Stamp. S. Tommaso d’ Aquino, 
Bologna. 

Paulucci, M. 1879. E'scursione scientifica nella Calabria 1877-78. Fauna malacologica. 
Specie terrestri e fluviatili enumerate e descritte. Coi Tipi dell’Arte della Stampa, 
Firenze. 

Salvadori, T. 1880. Ornitologia della Papuasia e delle Molucche. Parte prima. Mem. R. 
Acc. Sc. Torino (2) 33: I-XVI, 1-574. 

Salvadori, T. 1881. Ornitologia della Papuasia e delle Molucche. Parte seconda. Paravia, 
Torino. 

Salvadori, T. 1904. Note on Tanysiptera dea. Ibis (8) 4: 551-554. 

Salvadori, T. & D’Albertis, L. M. 1879. Catalogo degli uccelli raccolti da L. M. 
D’Albertis durante la seconda e terza esplorazione del Fiume Fly negli anni 1876 e 
1877. Ann. Mus. Civ. St. Nat. Genova 14: 21-147. 

Savi, P. 1827-1831. Ornitologia Toscana. Nistri, Pisa. 

Scopoli, G. A. 1763. Entomologia Carniolica exhibens insecta Carmioliae indigena et 
distributa in ordines, genera, species, varietates, methodo Linneana. Typis I. T. 
Trattner, Vindobonae. 

Scopoli, G. A. 1769. Annus I. Historico-Naturalis. Sumtib. Ch. G. Hilscheri, Lipsiae. 

Stresemann, E. 1975. Ornithology from Anitstotle to the Present. Harvard Univ. Press. 

Trischitta, A. 1950. Per una Nomenclatura Biologica ternaria plurinominale. Atti Acc. 
Peloritana det Pericolanti, Classe Scienze Fis. Mat. e Nat. 48: 1-40. 

Trischitta, A. 1952. Per Vauspicata discussione intorno alla nomenclatura biologica 
ternaria plurinominale di recente proposta. Boll. Zool. 11: 39-47. 

Trischitta, A. 1967. Systema Naturae usque ad genera viventium atque fossilium 
secundum nomenclaturam ternariam plurinominalem expositum. Atti Acc. 
Peloritana dei Pericolanti, Classe Scienze Fis. Mat. e Nat. 49: 263-274. 


Addresses: C. G. Violani FLS, Dipartimento di Biologia Animale, University of Pavia, 
Piazza Botta 9, 27100 Pavia, Italy. F. Barbagli FLS, Museo di Storia Naturale 
dell’ Universita, Sezione di Zoologia ‘“‘La Specola’’, Via Romana 17, 50125 Florence, 
Italy. 


© British Ornithologists’ Club 1997 


F. Barbagli et al. 145 Bull. B.O.C. 1997 117(2) 


Scopolhi, Linnaeus and the Wallcreeper 
Tichodroma muraria 


by Fabio Barbagli, Fausto Barbaglhi & Carlo Violani 


While examining letters written to Carl Linnaeus (Rashult 1707- 
Hammarby 1778) by Giovanni Antonio Scopoli (Cavalese 1723—Pavia 
1788) now preserved in the Linnean Society’s Library, London, and 
the respective replies published by Cobelli & Delaiti (1889) and 
recently by Soban (1995), we found interesting details on the 
nomenclature of some animals including the Edible Dormouse Myoxus 
glis (Violani & Zava 1995) and the Wallcreeper Tichodroma muraria. 
This bird was not included in the tenth edition of Systema Naturae 
(1758) probably because being a species foreign to Sweden it was 
unfamiliar to Linnaeus. Scopoli was a faithful correspondent of 
Linnaeus. He was employed by the Austrian Imperial Government as a 
physician to the quicksilver miners of Idria in Carniola (the region 
around Ljubljana in Slovenia) and his salary was supplemented by the 
tax money collected on wine sold in the area (Scopoli 1786-1788). 
Although isolated, Scopoli cultivated interests in many fields of the 
natural history of Carniola, botanizing, collecting insects and making 
observations on the local birds, during a long residence in Idria and on 
his trips through Slovenia and Friuli. | 

He wanted to send a specimen and communicate first-hand details of 
the Wallcreeper to Linnaeus so that his Swedish correspondent could 
include it in a future revised edition of Systema Naturae. In a long 
letter to Linnaeus, dated 28 January 1762, Scopoli announced that he 
had sent a Upupa muraria in a box together with other scientific 
material. However Linnaeus was unable to find the bird in the 
consignment when he opened the box, and so noted down on the left 
margin of Scopoli’s letter: ““Hanc non reperi in cistula; certe neglexit 
imponere Scopoli ...’’ [“‘I did not find this in the box; certainly Scopoli 
must have forgotten te: put iit ai.” 

On 11 February 1762, the Ttalian author apologized for his 
negligence and wrote: “‘... In cysta, pro Te, Cl. Gronovio missa, non 
invenies Picum murarium Aldrov _.. Nescio enim quo fato, apud me denuo 
remanserit, mittam tamen alia vice. Avis haec non est Picus, sed meo 
tudicio Upupa corpore supra cinereo, gula alba abdomine cauda alisq. 
nigris: bast remigibusq. primarius semirubris: tribus primis maculis duabus 
albis ...” [* ... In the box, sent to you through the celebrated 
Gronovius, you will not find the Picus murarius of Aldrovandi... I do 
not know by what mischance it remained with me, but I will send it 
some other way. This bird is not a woodpecker, but in my judgement 
an Hoopoe, with ash grey upperparts, throat white, belly, tail and wings 
black with the base of the primaries half-red, the three outer ones with 
two white spots... ’’]. 

At last, with a . long letter written by Scopoli on 7 April 1763, 
Linnaeus received a specimen of the bird accompanied by an accurate 
description of Merops murarius. Scopoli wrote: “‘... En descriptionem 


F.. Barbagli et al. 146 Bull. B.O.C. 1997 117(2) 


Gliris Romanorum et Pict muraru, quibus nonnullas alias observationes 


adiungo. ... Merops (murarwus), cinerascens, rostro abdomine cauda 
alisque nigris, remigibusque primaris semicoccinets, rectricibus apice 
albentibus ... Habitat et nidificat in Arcibus elatioribus et desertis, nec non 


> 


in turribus ...°’ [“‘Here is the description of the Dormouse of the 
Romans and of the Wallcreeper, to which I am adding some other 
observations ... Merops murarius, ashy coloured, bill, abdomen, tail 
and wings black, the primaries half scarlet, the rectrices white-tipped 
... It inhabits and nests on the higher solitary fortresses, as well as on 
towers 42)" |: 

A longer, detailed description followed. This was meticulous when 
giving body characters, and he obviously had a freshly killed specimen 
in winter plumage in front of him when writing the letter, since he 
was able to describe the shape of the nostrils, of the tongue 
and of the palate. A vivid report of the Wallcreeper’s behaviour 
revealed that Scopoli had observed the bird in life personally: ““Sub 
finem Autumni miegrat solitarius, volatu vago, remigante, muto. Aedificia 
elatiora, turres et Arces adit, muris insidet, super hos saltitando reptat, 
fenestras et latebras recognoscit, araneas devorat, muscas segniores capit, 
inquietus, frigoris amans, numquam pinguescens.’ |[‘“Towards the end 
of Autumn it migrates alone with a wandering, flapping, silent flight. It 
visits the higher buildings, towers and fortresses, sits on the walls, 
creeps on them hopping, explores windows and concealed places, 
devours spiders, catches the slower flies, restless, loving cold weather 
and never getting fat’’]. 

On 17 May 1763 an enthusiastic Linnaeus replied from Uppsala, 
gratefully acknowledging Scopoli’s helpful information: “‘Epistolam 
tuam V.A.d.7 aprilis data rite accepi, ... perplacuit pulcherrima avis 
europaea Picus muralis dicta...’ [“‘I have safely received your letter 
dated 7th April, ... I liked very much the beautiful European bird 
called Picus muralis ...’’]. 

Linnaeus included it as a new species under the name of Certhia 
muraria in his 12th edition of Systema Naturae (1766), where he 
acknowledged the information received. Later, Scopoli (1769) 
published a very similar diagnosis for Certhia murania. 

An interesting point concerns the type-locality of Certhia muraria 
Linnaeus 1766. Although briefly mentioning other authors, such as 
Gesner, Aldrovandi, Kramer and Brisson, Linnaeus clearly relied 
on Scopoli’s contribution for the printed description of the 
Wallcreeper. Since Scopoli’s specimen is stated to originate from 
Carniola, according to correspondence with Linnaeus, we believe that 
the Linnean type-locality (“Habitat in Europae australis turribus, 
arcibus’’) of Tichodroma muraria muraria (1...) can be restricted to 
Carniola in Slovenia. 


Acknowledgements 


The Authors wish to express their gratitude to Miss Gina Douglas, Librarian, and to the 
Linnean Society of London, for their kind permission to study and transcribe the 
Scopoli—Linnaeus correspondence, held in their care. 


D. Parkin 147 Bull. B.O.C. 1997 117(2) 


References: 

Cobelli, G. & Delaiti, C. 1889. Lettere inedite di Carlo Linneo a Giovanni Antonio Scopolzi. 
Tip. Roveretana, Rovereto. 

Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum Classes, Genera, 
Species cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima, 
Reformata. Laurentii Salvii, Holmiae. 

Linnaeus, C. 1766. Systema Naturae per Regna Tria Naturae, secundum Classes, Genera, 
Species cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Duodecima, 
Reformata. Laurentii Salvii, Holmiae. 

Scopoli, G. A. 1769. Annus I Historico-Naturalis. Sumtib. Ch. G. Hilscheri, Lipsiae. 

Scopoli, G. A. 1786-1788. Deliciae florae et faunae Insubricae. 3 vols. Typ. Reg. et Imp. 
Monasterii S. Salvatoris, Ticini. 

Soban, D. 1995. Linné’s letters to Scopoli 1761 — 1773. Prirodoslovno Drustvo Slovenije, 
Ljubljana. 

Violani, C. & Zava, B. 1995. Carolus Linnaeus and the Edible Dormouse. Hystrix (n. s.) 
6: 109-115. 


Addresses: Fabio Barbagli, Via Margaritone 20, 52100 Arezzo, Italy. Fausto Barbagli 
FLS, Museo di Storia Naturale dell’ Universita, Sezione di Zoologia “‘La Specola’’, 
Via Romana 17, 50125 Florence, Italy. C. G. Violani FLS, Dipartimento di Biologia 
Animale, University of Pavia, Piazza Botta 9, 27100 Pavia, Italy. 


© British Ornithologists’ Club 1997 


Molecular probes for identifications of raptors 


by D. Parkin 


Research into the Red Kite Milvus milvus at Nottingham resulted in the 
isolation of a clone DNA that is inherited in a sex-limited fashion. It 
reveals a multi-band profile that is transmitted more or less faithfully 
from mother to daughter. This ‘matrilineal’ pattern is typical for DNA 
that is sited on the female-specific (W) chromosome. Analysis of a 
series of unrelated female kites from Germany and Spain revealed that 
there were 20 different patterns among 27 nest sites. There was no 
evidence of a common pattern between the two regions. 

A long series of females from Wales revealed only two profiles, 
suggesting that this population is distinctly less variable. This finding is 
supported by the analysis of multi-locus DNA profiles in kites from 
these three regions. 

A single locus was analysed using an oligonucleotide probe. The 
number of alleles detected was significantly less in the birds from 
Wales, whereas those from Germany and Spain did not differ. 

All these results suggest that Red Kites from Wales are genetically 
depauperate. 

Interestingly, a southern isolate of the Welsh populations revealed a 
significant difference in genetic structure. First, the two matrilines 
differed in relative frequency, and second, the single locus data 
differed. The rarer of the matrilines was very similar to a German 
profile, suggesting the possibility that a bird from this region had 


D. A. Russell et al. 148 Bull. B.O.C. 1999-47 @} 


colonised South Wales at some time close to the date of spread from the 
traditional range in mid-Wales. 

In the late 1980s, it was decided to attempt to re-establish the Red 
Kite into an area of southern England from which it has been absent for 
over a hundred years. Blood samples were taken from the released birds 
which were also marked with rings and patagial (wing) tags. When 
breeding commenced, the identity of pairs was determined visually, 
and their success monitored. Blood samples taken from the nestlings 
permitted confirmation of identity and parentage. 

Although numbers were very small, evidence suggested that birds 
from Wales bred later and were less successful than the main release 
from Spain. This supports the hypothesis that the Welsh birds are 
not only less variable (inbred) but also less successful (inbreeding 
depression). Supporting evidence will be presented, and recommen- 
dations made for future releases in any re-introduction programme. 


Address: D. Parkin, Department of Genetics, School of Medicine, Queen’s Medical 
Centre (University of Nottingham), Nottingham NG7 2UH, U.K. 


© British Ornithologists’ Club 1997 


X-raying the Gods: what were the mummified 
Horus falcons of Egypt? 


by D..A. Russell, B. Galeb & R. Hoath 


As part of the Egyptian Exploration Society (U.K.) investigation of the 
social and temporal context of the Sacred Animal Necropolis at 
Saqqara, Egypt, in 1992, 1994 and 1995, identification to species level 
was attempted for some 200 mummified ‘falcons’. ‘These were amongst 
the hundreds of thousands deposited in underground galleries from 
c. 600BC to 100AD by devotees of the religious cult of Horus worship. 
Many of the remains were in poor condition, partly as a result of the 
‘hot dipping’ method of mummification used and partly because of the 
age and condition of the material at the time of its mummification 
(many specimens were partial skeletons, mixed species or other 
material, e.g. eggshells, twigs or shrews). 

Identification by manual unwrapping was both laborious and 
destructive, owing to the friable nature of the material, and the 
wrapping of some specimens was too fine to permit destructive 
sampling. Consequently, a sub-sample of mummies was X-rayed on 
site. A portable army field machine, manufactured by the Massiot 
Society, was operated at 10mA and 60kV by Drs R. and M. 
Lichtenberg with the kind permission of the Mission Archéologique 
Francaise du Bubasteion. Apart from the need for accurate scaling, 


K. E. Mikhailov 149 Bull. B.O.C. 1997 117(2) 


most of the difficulties in resolution of the bones turned out, on 
subsequent unwrapping, to be unresolvable problems related to the 
charred, broken or non-skeletal nature of the material. Determinations 
were made mostly on the basis of long-bone (especially tarsal) and skull 
measurements, as these showed reduced dependency on the precise 
conformation of the bird within the mummy, which was often not as 
indicated by the external wrapping. Simple proportionality indices 
were utilised where there were gender overlaps in size range between 
related species. 

Perhaps surprisingly, considering the enormous numbers of birds 
mummified at the site, most identified remains were of birds of prey 
(both residents and passage migrants) likely to have been common in 
the immediate area. Most were Kestrels Falco tinnunculus, Lanner 
F.. biarmicus and Barbary Falcons F. pelegrinoides, though species of 
hawks, harriers Circus spp., eagles, vultures and some other species of 
falcon were also found. However, species only utilising the migration 
pathway along the adjacent Red Sea coast (e.g. Lesser Spotted Eagle 
Aquila pomarina) were absent, as were the expected three species of 
kite. The latter, including the falcon-like Black-shouldered Kite Elanus 
caeruleus, must have been common in the area at the time and have 
been recorded from other archaeological sites in Egypt. 

Allied with the burial of some individuals with appropriate 
‘food-parcels’ (including a Marsh Harrier Circus aeruginosus with a 
small mummified snake), the mummies show that the ancestral 
inhabitants of the Nile valley had considerable, detailed, knowledge of 
the birds of prey of the region. A wide range of birds of prey (but 
possibly not all) appear to have been sacred to Horus, the sky God, son 
of Osiris, with whom the living King was closely identified. 


Addresses: D. A. Russell, American University in Cairo, PO Box 2511, Cairo, Egypt. 
Present address: Pest Management Department, Natural Resources Institute, 
University of Greenwich, Chatham Maritime, Chatham, Kent ME44TB, U.K. B. 
Galeb, Egypt Exploration Society, 3 Doughty Mews, London WC1, U.K. R. Hoath, 
American University in Cairo, PO Box 2511, Cairo, Egypt. 


© British Ornithologists’ Club 1997 


Bird taxonomy based on eggshell structure 


by K. E. Mikhailov 


In terms of evolution, the taxon is the result of a particular 
morphogenetic trajectory: it is an objective category which is 
independent of phylogenetic (=genealogical) hypotheses and logically 
precedes them. Major distinctions between taxa, such as eggshell 
structures, are very important since they enable the recognition of 


K. E. Mikhailov 150 Bull. B.O.C. 1997 117(2) 


different trajectories amongst a mosaic of similarities. A working 
assumption is that two families of birds within the same order should 
not exhibit different eggshell structures, since such differences indicate 
substantial morphogenetic separation. It is also assumed that 
similarities in the basic eggshell structure suggest that the families 
being compared are members of a single evolutionary trajectory. 

Comparisons of eggshell structures have led to the following 
conclusions. Ratite, galliform and anseriform birds show ratite-like 
eggshell structures; therefore each of these groups is taxonomically very 
distinct from the neognathe bulk of families. The casuariiform birds 
(emu and cassowaries) are separate within the traditional paleognathes, 
and the Cracoidea (Megapodiidae and Cracidae) are separate from the 
other galliform birds. Three orders of aboreal neognathes, namely 
Cuculiformes, Coraciiformes and Piciformes, also exhibit somewhat 
ratite-like eggshell structures. A group consisting of the Psittaciformes, 
Strigiformes and Caprimulgiformes, and another including the 
Apodiformes, Coliiformes and Trogoniformes, form separate structural 
assemblages of arboreal orders, as do the Passeriformes. All gruiforms 
and charadriiforms (amongst which the jacanas are most distinct) have 
generally similar eggshell structures. ‘The unusual suborder Ciconii 
sensu Sibley and Ahlquist is an assemblage of high ranking taxa, 
including ‘the Pelecaniformes (which may be separate from the 
Phaethontes, but allied with the Balaenicipitidae) with the Ardeiformes 
(Ardeidae, Threskiornithidae, Scopidae), Falconiformes, Accipitri- 
formes (Accipitridae+ Pandionidae), Ciconiiformes (Cicontidae only), 
Procellariiformes, Gaviuformes, Sphenisciformes, Podicipediformes 
and Phoenicopteriformes as very separate groups. The Dromaiidae+ 
Casuariidae, Accipitridae+ Pandionidae, Ciconiidae and Bucerotidae 
(possibly all Upupiformes) have eggs with the most aberrant eggshell 
structures among avian families. 


Acknowledgement 


- Support from PalSIRP grant-1995 and RFBR grant N 96-04-50822 is gratefully 
acknowledged. 


Address: K. E. Mikhailov, Paleontological Institute of the Russian Academy of Sciences, 
Profsojuznaja St. 123, Moscow 117647, Russia. 


© British Ornithologists’ Club 1997 


Books Received 151 Bull. B.O.C.A99% 1172) 
BOOKS RECEIVED 


del Hoyo, J., Elliot, A. & Sargatal, J. (eds) 1996. Handbook of the Birds of the World. Vol. 
3, Hoatzin to Auks. Pp. 752, 60 colour plates, 389 colour photographs, 577 distribution 
maps, c. 8000 bibliographical references. Lynx Edicions, Barcelona. ISBN 84-87334- 
20-2. c. £105 (plus p. & p.). 31 X 24cm. 


This superbly illustrated volume is the third part of the HBW series now structured to 
be published in 12 volumes, subsequent parts planned to be published every 18 months 
and eventually at shorter intervals. With publication of Vol. 3, the first 69 families of 
birds have been dealt with out of the envisaged world total of 176, in less than 4 years. 
HBW benefits from having been supported by BirdLife International from the very early 
days, with the provision of up to date information on the conservation. status of many 
species. 

The foreword and frontispiece by Robert Bateman are followed by a relatively short 
4-page introduction. Readers are referred to the detailed general introduction to the 
series in Vol. 1 (pp. 15-33), the guiding taxonomic principles of which are summarised 
briefly as fairly conservative at the macrosystematic level, increasingly flexible at lower 
levels, and subspecifically aiming to follow latest developments appearing to have gained 
reasonable acceptance. Vol. 3 is stated, however, to probably represent the single most 
significant taxonomic deviation from traditional usage in the series, e.g. the Plains- 
Wanderer Pedionomus torquatus, endemic to Australia, is placed in Charadriiformes; the 
taxonomically controversial Hoatzin Opisthocomus hoazin is placed in the monotypic 
order Opisthocomiformes; the sandgrouse (Pteroclidae) have been awarded a separate 
order, Pterocliformes. Mention is also made of the new IUCN criteria for threat (the 
Mace-Lande categories being no longer used): Vulnerable, Endangered and Critically 
Endangered. 

The family and species accounts generally retain the style, layout and headings of the 
first two volumes of the series, the family accounts continuing to be illustrated by a 
superb range of colour photographs. One significant innovation is the inclusion of notes 
on voice in the Descriptive Notes section (originally intended only for the passerines) for 
all species in the Gruidae and Rallidae. An example of the level of detailed information 
given is, for Status and Conservation of the Rallidae, the list of rail taxa (including 
subspecies) extinct since 1600 linked to the high incidence of flightlessness, their status as 
island endemics and the probability of their having succumbed to predators introduced 
by man. 

Species names are also given in French, German and Spanish (publication of 
recommended Spanish names continues elsewhere) in addition to the scientific and 
adopted English names. Several examples of more recent developments affecting 
taxonomy and terminology retained by Vol.3 are: Turnix sylvatica (9 subspecies 
recognised) rendered as Common Buttonquail but in the other 3 European languages 
more nostalgically as Turnix d’Andelousie/Laufhtihnchen/Torillo Andaluz; Haematopus 
ostralegus becomes Eurasian Oystercatcher/Huitrier pie/Austernfischer/Ostrero euro- 
asiatico; Catharacta antarctica (3 ssp.) becomes Brown Skua with C. skua separated as 
monotypic; for the Laridae, the so-called white-headed gulls group is identified as having 
proved extremely challenging (species limits among species pairs in the Herring Gull 
complex being controversial). Larus argeniatus (4 ssp.) is retained as Herring Gull/ 
Goéland argenté/Silberm6we/Gaviota argéntea: L. cachinnans (5 ssp.) becomes Yellow- 
legged Gull/Goéland leucophée/WeiBkopfm6we/Gaviota patiamarilla: L. armenicus 
(monotypic) becomes Armenian Gull/Goéland d’Arménie/Armenienméwe/Gaviota 
Armenia; L. fuscus is allocated 4 ssp. including heuglini. 

The list of the 60 beautifully reproduced colour plates illustrating plumage 
characteristics (including most subspecies) credits the individual artists, the complete 
international Vol. 3 team being Norman Arlott, Hilary Burn, Angels Jutglar, Francesc 
Jutglar, lan Lewington, Chris Rose, Lluis Sanz, Etal Vilaré and Ian Willis. The plates 
also incorporate a simple graduated scale allowing size to be appreciated. Vol. 3 ends with 
a list of references to the original scientific descriptions, a general list of all references and 
an index. 

Vol. 3, as the previous two volumes, is such excellent value for money (particularly if 
special publication offers are taken up) that it is highly recommended for all devotees 
from the enthusiastic amateur to the dedicated professional. 

S.J.F 


Books Received 1 ey Bull. B.O.C. 1997 147(2) 


Feduccia, A. 1996. The Origin and Evolution of Birds. Pp. x+420, numerous text-figures. 
ISBN 0-300-06460-8. Yale University Press. £45.00. 28.5 x 21.5 cm. 


Tremendous advances have been made in knowledge of avian evolution in the last 20 
years, from the study of new fossil material. Much of it has appeared in publications not 
readily available to general ornithologists. A broad, up to date survey by a leading 
researcher in the field thus fills a major gap in the bird literature. Indeed, keeping 
completely up to date in a book of this scope is scarcely possible; Feduccia was not able 
to provide a full account of important Jurassic and Cretaceous fossils that were still under 
study at the time of his writing. 

A great merit of this book is that the necessary technical details are made intelligible to 
a non-specialist, and for the most part the whole text is highly readable. Considerations 
not only of zoogeography and continental movements, but also of ecology and behaviour, 
are brought to bear on the task of presenting a historical and functional interpretation 
of the whole course of avian evolution. Readers unfamiliar with what has been revealed 
by recent research will be amazed by the extraordinary diversity and succession of 
independently evolved adaptive types over tens of millions of years. Tentative 
interpretation of avian evolution on the basis of present-day distributions turns out to be 
almost wholly fallacious. Families now endemic to a continent and so thought to have 
evolved there, for example colies in Africa, were once widespread. Early forms of todies 
have been found in North America and France, showing that the five species in the West 
Indies are a surviving remnant and throw no light on the origin of the family. An 
overwhelming impression, from Feduccia’s survey, is that our present world avifauna, 
judged by body size and structure alone, is rather a diminutive and impoverished one by 
comparison with what there once was; but there has been compensation in the recent 
proliferation of the passerines, which may have reached a level of sophistication of 
behaviour in advance of anything that preceded them. 

The illustrations are excellent, adding greatly to one’s appreciation of the text. The 
latter is slightly marred by minor errors and misprints that seem to be due to inadequate 
proof-reading. Further, some of the interpretation, especially concerning the very early 
evolution of birds, is controversial, and this will not always be apparent to the 
non-specialist reader; but this hardly detracts from an overall judgment that this is one of 
the most interesting books on birds to have appeared for years. 


Woods, Robin W. and Anne. 1997. Atlas of Breeding Birds of the Falkland Islands. 
Pp. 190, 73 maps. Anthony Nelson Publishers, in association with Falklands 
Conservation. ISBN.0-904-614-60-3. £25.00. 25 x 17 cm. 


This is the first breeding bird atlas published for a South American country, and shows 
the importance of the Falkland archipelago for thriving populations of species that have 
declined dramatically in southern South America. The authors have 40 years personal 
knowledge of these islands, and this book is the outcome of contributions from about 160 
observers, who sent in survey forms over a period of ten breeding seasons between 
1983/84 and 1992/93. The delightful line drawings for each species are by Staff Sergeant 
Geoffrey McMullan. 

This work is a classic example of how the observations of a wide range of birdwatchers, 
both professional and amateur, can be combined to produce a reference of real value. 
About 80 (51%) of these were Falkland Islanders or contract workers; 47 (30%) were 
military personnel, temporarily stationed on the islands, or visiting in RN/Royal Fleet 
Auxiliary ships; 29 (19%) were visiting tourists to the Falklands. 

This is an important book, and with continuing exploitation of fishery resources, and 
the threatened development of off-shore oil now imminent, its publication is very timely. 
The excellent maps provide unique distributional data of all established breeding species 
and, more importantly, show where current knowledge is incomplete, and where further 
surveys are necessary. The Atlas will be of interest to Falkland Islanders and will be a key 
reference for future visitors and tourists, because there is much work still to be done. It 
will also be of interest to ornithologists worldwide, to anyone studying island faunas, and 
to people who are concerned about the environment. Maps included show the 
distribution of introduced mammals, including domestic/feral Cat, Common and Black 
Rat and European Rabbit. 

The result is well-produced, and compares very favourably with the previous work 
from the same stable, Robin Woods’ Guide to Birds of the Falkland Islands ae hs 


NOTICE TO CONTRIBUTORS 

Papers, from Club Members or non-members, should be sent to the Editor, Dr 
D. W. Snow, The Old Forge, Wingrave, Aylesbury, Bucks HP22 4PD, U.K., 
and must be offered solely to the Bulletin. After 15 September 1997, they 
should be sent to Dr C. J. Feare, 2 North View Cottages, Grayswood 
Common, Haslemere, Surrey GU27 2DN. They should be typed on one 
side of the paper, with treble-spacing and a wide margin, and submitted in 
duplicate. The style and lay-out should conform with usage in this or recent 
issues of the Bulletin. 


A contributor is entitled to 10 free offprints (16 if 2 or more authors) of the pages of the 
Bulletin in which his contribution, if one page or more in length, appears. Additional 
offprints or offprints of contributions of less than one page may be ordered when the 
manuscript is submitted and will be charged for. Authors may be charged for proof 
corrections for which they are responsible. 


MEMBERSHIP 
Only Members of the British Ornithologists’ Union are eligible to join the 
Club, and to receive (postage free) four quarterly issues of the Bulletin, and the 
annual index, for an annual subscription of £12 (or U.S. $26). Applications, 
enclosing the annual subscription, should be made to the Hon. Secretary 
(address as below). 


NON-MEMBER SUBSCRIBERS & APPLICATIONS FOR BACK 
NUMBERS OR OTHER PUBLICATIONS 

The Bulletin (for 1997 onwards), together with annual index, may be 
purchased (postage free) by non-members on payment of an annual sub- 
scription of £22 (or U.S. $45) on application to The Publications Officer, S. J. 
Farnsworth, Hammerkop, Frogmill, Hurley, Maidenhead, Berks SL6 5NL, 
U.K. Single issues, and back numbers of the Bulletin, and also books in the 
BOC Occasional Publications series may similarly be obtained, on request 
to him. 


PAYMENTS 

All amounts quoted are net and should be paid in £ sterling, if possible. 
Payments in other currencies must include a further £4 for UK bank charges 
(except for annual rates in U.S. dollars, which are inclusive). All cheques or 
drafts should be made payable to the British Ornithologists’ Club. If preferred, 
remittances may be made by bank transfer direct to the Club’s bank 
account—Barclays Prime Account, Dale House, Wavertree Boulevard, 
Liverpool L7 9PQ, U.K. (Sort Code 20-00-87 Account No. 10211540), with 
confirmation to the Hon. Treasurer, D. J. Montier, Eyebrook, Oldfield Road, 
Bickley, Bromley, Kent BR1 2LF. 


CORRESPONDENCE 
Correspondence on membership, changes of address and all other matters 
should be addressed to the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA, U.K. For details 
of Club Meetings see inside front cover. 


Registered Charity No. 279583 


CONTENTS 


Page 
CLUB NOTICES.) Meetings: 2a ON see ine A ee Car 81 
Avian Taxonomy from Linnaeus to DNA 
CHERE ROA. Preface! od 00), fa Akasa nec 83 
GREENWOOD, J. J. D. Introduction: the diversity of taxonomies ....... 85 
ZINK, R: M. Species concepts. )...0 5 4 body ean se ll yr 
SNOW, D. W. Should the biological be superseded by the phylogenetic 
SPECIeS CONCEPE! ). 6 scgie oni aw Aes 110 
COLLAR, N. J. ‘Taxonomy and conservation: chicken and egg......... 122 
VIOLANI, C. G. & BARBAGLI, F. Salvadori, Giglioli and Arrigoni: aspects 
of the trinomial controversy in Italian ornithology.............. 137 
BARBAGLI, F., BARBAGLI, F. & VIOLANI, C. Scopoli, Linnaeus and the 
Wallcreeper Tichodroma muraria.. 0.6... bss ee 145 
PARKIN, D. Molecular probes for identification of raptors ........... 147 
RUSSELL, D. A., GALEB, B. & HOATH, R. X-raying the Gods: what were the 
mummified Horus falcons of Egypt?. .\...........4)) er 148 
MIKHAILOV, K. E. Bird taxonomy based on eggshell structure ........ 149 
BOOKS RECEIVED 2.3.05 6.0.c8 6 wig eo cee ose acai pln aie we) dl nen 151 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 
mail, after their current subscription has been paid. 


COMMITTEE 


D. Griffin (Chairman) (1993) 


Miss H. Baker (1994) 
Revd T. W. Gladwin (Vice-Chairman) (1993) S 


J. Farnsworth (1990) 


D. J. Montier (A/Hon. Treasurer) (1996) R. E. F. Peal (1993) 
Cdr M. B. Casement, OBE, RN Dr R. P. Prys-Jones (1995) 
(Hon. Secretary) (1996) N. H. F. Stone (1995) 


Hon. Editor: Dr D. W. Snow 
Chairman of Publications Sub-committee: Mrs A. M. Moore 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


(oT | 
BSGS ISSN 0007-1595 
3 |\RD 


Bulletin of the 


British Ornithologists’ Club 


Edited by 
Dr D. W. SNOW 


NOW 1 ¢ GU / 
Ue U 0 1771 


LIBRARIES 


Volume 117 No. 3 September 1997 


FORTHCOMING MEETINGS 


Advance notice of meeting dates for 1998. Nine meetings are being arranged 
for the following Tuesdays: 20 January (Ian Burrows on Birds of Papua New 
Guinea—see below), 17 March (Tony Prater on Waders), 21 April (Dr W. R. P. 
(Bill) Bourne on Birds and Islands), 19 May (AGM and social evening—with 
informal “‘mini-talks” as in 1997), 14 July, 15 September, 13 October and 17 
November. Details of speakers on these dates will be published when finalised. 


Tuesday 20 January 1998. Dr Ian Burrows B.Sc., Ph.D. will speak on ‘*The 
Birds of Papua New Guinea’’. Ian has been a keen birder for many years, with a 
passion for rare and elusive species. After spending a year as a Warden of Cape 
Clear Bird Observatory in 1975, he obtained a Ph.D. in Applied Microbiology from 
Aston University in 1980. He has spent ten years (1986-1995) in Papua New Guinea 
lecturing in Microbiology, Ecology and Biology at the University of Papua New 
Guinea, in Port Moresby. He has visited and watched birds in every province of 
PNG, apart from Bougainville, and has conducted extensive research and 
consultancy programmes on the Melanesian Scrubfowl, Macgregor’s Bird of 
Paradise, and a wide range of rainforest birds. A fluent pidgin speaker, he has 
worked as an in-field consultant for the British and Australian Broadcasting 
Corporations and the World Wildlife Fund. In 1997, together with Phil Gregory, he 
has founded Sicklebill Safaris Ltd., specialising in tours to Melanesia and Australia 
in search of rare and little known species.. 

Applications to the Hon. Secretary by 4 January, please. 


Tuesday 4 November 1997. Tony Marr will speak on “An Atlantic Seabird 
Odyssey’”’. Tony has had a life-long interest in seabirds, especially through 
‘pelagics’ in the eastern Atlantic, and he has pursued them from a wide variety of 
boats and craft, off Portugal, The Canaries, Madeira, West Africa, and the South 
Atlantic. Having retired early to devote more time to birds, he is well-known to 
many as a tour leader to many parts of the world, and as a lecturer and author of 
articles on seabird topics. He serves on the BOU Records Committee, and the 
Seabirds Advisory Panel for British Birds. 

Applications to the Hon. Secretary by 21 October (Trafalgar Day!), please. 


Meetings are held in the Sherfield Building of Imperial College, South 
Kensington, London SW7, at 6.15 p.m. for 7 p.m. The nearest Tube station is at 
South Kensington, and car parking facilities are available; a map of the area will be 
sent to members, on request. The cash bar is open from 6.15, and a buffet supper, of 
two courses followed by coffee, is served from about 7.00. (A vegetarian menu can be 
arranged if ordered at the time of booking.) Informal talks are given on completion. 


Overseas Members visiting Britain are particularly welcome at meetings. 
For details in advance, please contact the Hon. Secretary, Cdr M. B. 
Casement, OBE, RN: Dene Cottage, West Harting, Petersfield, Hants 
GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1997 


Apart from single copies made for the purposes of research or private study, or 
criticism or review, as permitted under UK law, no part of this publication may be 
reproduced, stored or transmitted in any form or by any means, except with prior 
permission in writing of the publishers, or in accordance with the terms of licences 
issued by the Copyright Licensing Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; 
for address see inside back cover. 


153 Bull. B.O.C. 1997417(8) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 117 No. 3 Published 19 September 1997 


ANNUAL GENERAL MEETING 


The Annual General Meeting of the British Ornithologists’ Club was 
held in the Ante-room of the Sherfield Building, Imperial College, 
London SW7 on Tuesday 20 May 1997 at 6 p.m. with Mr D. Griffin in 
the Chair. Apologies had been received from R. E. F. Peal, S. J. 
Farnsworth, Professor C. J. Feare and Mrs Mary Muller. 23 Members 
were present. 

The Minutes of the Annual General Meeting held on 21 May 1996, 
which had been published (Bull. Brit. Orn. Cl. 116: 201-202), were 
approved and signed by the Chairman. 


The Report of the Committee for 1996, which had been published 
(Bull. Brit. Orn. Cl. 117: 1-3), was presented. The Chairman drew 
attention to the highlights: eight evening meetings, with an average 
attendance of 36; the highly successful joint meeting with the Linnean 
Society, in March, organised by Dr R. A. Cheke; the very enjoyable 
excursion to Gilbert White’s house at Selborne on 31 May; and the 
publication of Vol. 116 of the Bulletin, containing a wide range of 
material, including colour plates of two new species. It is hoped that 
these colour plates will become a regular feature, and the Club is 
indebted to Tom Gladwin for arranging future sponsorship. 
Membership had increased to 567 (550)—-322 U.K. and 245 overseas. 

1996 also saw the publication of the second in the Occasional 
Publication series—Manuscripts and Drawings in the Ornithology and 
Rothschild Libraries of the Natural History Museum at Tring, by Mrs 
F. E. Warr. Several other publications were in the pipeline, and the 
Chairman thanked Mrs A. M. Moore and the Publications 
Sub-committee for all their work. Approval of the Report was proposed 
by Mrs A. M. Moore, seconded by Iain Bishop, and approved by all 
present. 

The Chairman then thanked the Hon. Secretary for the arrange- 
ments for the evening meetings, and also for the additional tasks he had 
taken on, including the painful learning of the intricacies of 
word-processing, the updating of address lists, and the coordination of 
the handover of the Treasurer’s tasks from John Farnsworth to David 
Montier. All Membership matters and subscription reminders were 
now handled by the Hon. Secretary, ably assisted by Helen Baker. 


The Accounts for 1996 were presented by David Montier, who had 
taken over as Acting Hon. Treasurer, on the resignation of John 
Farnsworth, as from November 1996. The accounts covered the year 
ended 31 December 1996, and included a statement of Committee 


Meetings 154 Bull. B.O.C. 1997 tiga 


Members’ Responsibilities, and Independent Examiner’s Report 
(Donald Reid & Co); copies of the accounts were distributed to all 
Members present. 

David Montier drew attention to the main points. The Income & 
Expenditure Account for 1996 showed a surplus of £5,959 (£5,549 in 
1995). Investment income included (with Income Tax recovered) a 
total of £9,912 from the Herbert Stevens Fund. £3,572 had been 
received from Sales of Publications, including £1,858 for sale of 
Bulletin back-numbers, much of which was from a single big order. 
Publication costs of the Bulletin were £10,118 (£8,526), the 1996 figure 
including £1,382 for the additional cost of colour plates. The cost of 
evening meetings was £3,454, whereas the income was £3,131, which 
represented a loss of £323 (£444 in 1995), but this figure would 
increase in 1997, due to the additional room hire charge of £120 per 
evening, as from December 1996. In general, the figures showed a fairly 
healthy state. 

Details of the Herbert Stevens Trust Fund were given on a 
separate statement, showing an end of year balance of £172,575 
(£162,791), and the investment portfolio a valuation increase of 
£10,886, over the year. 

In answer to a question, Mr Montier replied that he would look into 
the suggestion that, since the 21 year term had now expired, the 
Barrington Trust Fund (valued at £577) should now be transferred to 
the General Fund, or possibly added to the (James Monk) Publication 
Fund. The Bird Atlas of Uganda Fund (£2,597) represented an 
advance sponsorship donation from Shell Uganda, and was being held 
in a separate deposit account, until the future costs and sponsorship 
plans were clarified. 

Dr James Monk warmly congratulated David Montier on the clarity 
and timeliness of these accounts. Adoption of the Accounts was 
proposed by the Acting Hon. Treasurer, seconded by Stan Howe, and 
approved by all present. 

The Chairman thanked David Montier for allowing himself to be 
persuaded to take on the task as Hon. Treasurer, and for presenting the 
audited accounts for this meeting. There had been many meetings 
between him and John Farnsworth during the past year, and the Club 
was very grateful to them both for achieving a smooth hand-over of 
responsibilities. David Montier, in response, paid tribute to the 
immense volume of work achieved by his predecessor during his term 
of office. Those tasks were now shared between three people. 

The Chairman expressed the thanks of all to the Trustees of the 
Herbert Stevens Trust Fund, for their management of these funds 
during the year, and also to Dr James Monk for his further donations to 
the Publications Fund. 


The Bulletin. Dr David Snow reported that the next issue, 117(2), 
to appear in June, would include five of the main papers, and several 
shorter papers presented at the joint meeting with the Linnean Society 
last year, which should attract wide interest. He had agreed with Chris 
Feare to hand over the Editorship in the course of September, and an 


Meetings 155 Bull. B.O.C. 1997 117(3) 


appropriate notice to contributors would be included inside the rear 
cover of 117(2). There was a good supply of contributions sufficient to 
the end of this year, and into 1998, and he was pleased with the policy 
to include regular colour plates. Dr Robert Prys-Jones said it was 
important to continue this, and Tom Gladwin saw no problem in 
finding the sponsorship necessary for the future. 

The Chairman warmly thanked Dr Snow, on behalf of the whole 
Club, for another excellent volume last year, and for his wise 
Editorship since 1991. He was grateful for his agreement to continue 
until Chris Feare was available to take over, in September. Thanks 
were also due to Mrs Mary Muller for her work in again producing the 
Annual Index for 1996, which would appear in June. 


Election of Officers and Committee. As retiring Chairman, and 
before proceeding to the election of Officers, David Griffin expressed 
his personal thanks to all those who had so readily helped in the 
running of the Club during his four-year term of office. In addition to 
those already mentioned, special thanks were due to Ronald Peal, 
who had been one of the leading lights of the Club ever since he 
became Hon. Secretary in 1971, and had been Chairman in the 
Club’s Centenary Year. In the past four years as Committee Member, 
we had all benefited greatly from his wealth of experience. Francis 
Stone, who continues on the Committee, is very active in sorting and 
cataloguing the Club’s archives. Effie Warr, who is the author of this 
years’s Occasional Publication No. 2, has for many years stored and 
managed our stock of Bulletin back-numbers. Professor Richard 
Chandler had kindly looked after our interests, and stored our 
equipment in Imperial College. Thanks are also due to Ron Kettle 
and Pat Sellar for organising the projector and audio equipment at our 
meetings. 

The Committee’s proposals had been published in Bulletin 117(1): 3, 
and no other nominations had been received. He proposed that Michael 
Casement be re-elected as Hon. Secretary, and David Montier be 
confirmed as Hon. Treasurer. This proposal was seconded by Dr James 
Monk and agreed by all present. 

The Committee proposed that Dr David Snow be re-appointed as 
Editor, until September, and Professor Chris Feare as Editor, on a date 
to be mutually agreed, Iain Bishop seconded this proposal, which was 
agreed by all present. 

The Committee proposed the election of Mrs A. M. Moore as 
Vice-Chairman, to replace Tom Gladwin, and is very glad that she has 
accepted this nomination, in view of her long-standing service to the 
Club, as Secretary 1989-95, and since as Chairman of the Publications 
Committee. Ron Kettle seconded this proposal, which was agreed 
unanimously. 

David Griffin said that his final pleasant duty, as retiring Chairman, 
was to propose the election of the Reverend T. W. Gladwin to succeed 
himself as Chairman. Tom had been a tremendous support during his 
own term of office, and had been very active in encouraging new 
activities for the Club. He had inspired and organised the two visits by 


Meetings 156 Bull. B.O.C. 1997 117(3) 


members to the Natural History Museum at Tring, and the visit last 
year to Selborne. He had also arranged sponsorship for the colour 
plates now appearing regularly in the Bulletin. This proposal was 
seconded by Stan Howe and agreed unanimously. 

‘The Revered ‘Tom Gladwin was declared elected as Chairman, and 
he received the good wishes of all, for his term of office. In response, he 
thanked David Griffin for his Chairmanship of the Club, throughout a 
period of considerable change, and for his encouragement for new ideas 
and re-organisation of Club business. He was especially pleased that 
the Committee could count on his continued support and advice, as a 
member of the Committee. 

He proposed that N. J. Redman be elected to fill the vacancy on the 
Committee, on the retirement of R. E. F. Peal. This was seconded by 
Iain Bishop, and agreed by all. | 


No other items for discussion had been notified in accordance with 
Rule (12), and meeting closed at 6.30 p.m. 


The eight hundred and sixty-seventh meeting of the Club was held on Tuesday, 8 April 
1997, at 6.15 p.m., in the Senior Common Room at Imperial College. 31 Members and 8 
guests attended. 

Members present were: D. GRIFFIN (Chairman), Dr C. J. HazEvort (Speaker), Miss H. 
BaKER, J. W. BARRINGTON, P. J. BELMAN, Dr W. R. P. BourRNE, Mrs D. M. BraDLey, 
D. R. CaLper, Cdr M. B. CasEMENT RN, Professor R. J. CHANDLER, Dr R. A. CHEKE, 
D. J. Fisher, Dr L. D. C. FIsHpooL, Rev. T. W. GLapwin, A. Gipss, J. A. JOBLING, 
C. A. R. HeEtm, R. M. Kettie, N. S. MaLtcotm, Dr C. F. Mann, D. J. MontTiser, R. G. 
Morcan, Mrs M. N. Mutter, R. E. F. Peat, Dr R. P. PrYs-jongs, S. J. R. RuMsEy, 
R. E. Scott, P. J. SELLAR, Dr D. W. Snow, N. H. F. SToNngE, C. W. R. STOREY. 

Guests attending were: M Brapiey, Mrs J. B. CaLprer, Mrs C. R. CasEMENT, Mrs 
J. M. Grapwin, Mrs S. Grirrin, A. Martin, Mrs M. Montier, R. RANrFrT. 

After dinner, Dr Cornelius Hazevoet gave a presentation, illustrated with colour slides, 
on ““The Birds of the Cape Verde Islands’’. 

After shorter visits in 1986-1987, the speaker was semi-resident in the Cape Verdes 
during the years 1988-1996 as project co-ordinator of the National Parks and Protected 
Areas Programme (NPPAP), an initiative carried out by the Instituto Nacionale de 
Investigagao e Desenvolvimento Agrario of Cape Verde and the Institute of Systematics 
and Population Biology (Zoological Museum) of Amsterdam. 

The Cape Verde Islands are an oceanic archipelago situated c. 500 km west of Senegal. 
There are ten islands (of which nine are inhabited) and a number of uninhabited islets. 
Situated at the latitudes of the Sahel zone, the climate is generally arid with unpredictable 
and irregular rains. Among the landbirds, desert types (such as larks of the genera 
Alaemon, Ammomanes and Eremopierix, and Cream-coloured Courser Cursorius cursor) 
take a conspicuous place in the avifauna. More significantly, there are several taxa 
endemic to the islands, many of which appear to be relics from Pleistocene periods, when 
the climate in both the western Sahara and the adjacent Cape Verde Islands was 
considerably more humid than today. 

While the climate deteriorated and taxa disappeared on the mainland, circumstances 
in the Cape Verde Islands remained relatively favourable for a longer period, enabling 
taxa to survive there, in isolation. These include a heron Ardea bournei, kite 
Milvus fascticauda, buzzard Buteo bannermani, and kestrels Falco neglectus and 
F. alexandri. Endemic taxa with a more problematic zoogeographical history include a 
peregrine F. madens, barn owl Tyto detorta, the enigmatic Raso Lark Alauda razae, 
Cane Warbler Acrocephalus brevipennis and sparrow Passer iagoensis. Although some of 
these are widespread (T. detorta) or abundant (P. iagoensis) in the islands, many of the 
endemics are extremely rare and seriously threatened. For instance, the total population 
of A. bournei does not exceed some 20 pairs (and perhaps less) in two colonies, each 
situated in a single tree, while numbers of M. fasciicauda are decreasing at an alarming 


Meetings 157 Bull. B.O.C. 1997 117(3) 


rate. The latter was once widespread in at least five of the islands, but today dwindling 
numbers only survive on the island of Santo Antao, and it will be extinct if no effective 
action is undertaken for its preservation. In 1988-92, the population of A. razae was 
estimated at c. 250 birds but recent visits could locate considerably less. Although this 
may partly be due to natural fluctuations in population size, this breeding taxon is 
extremely vulnerable to ground predators (cats, rats and dogs) which may be 
inadvertently released in its only resort (uninhabited Raso islet; 7 km?) by visiting 
fishermen. The Cane Warbler A. brevipennis once occurred on three islands but is now 
only found in a patchy distribution on Santiago, with a population not exceeding some 
500 pairs. 

The situation for the seabirds is equally, it not even more, drastic; numbers have been 
greatly reduced due to the constant and centuries-long persecution by local fishermen, 
who harvest eggs and young for food, and destroy adults for no particular purpose. 
Populations of Red-billed Tropicbird Phaethon aethereus and Brown Booby Sula 
leucogaster are estimated to have declined from c. 1000 and 10,000 during the late 19th 
century to c. 100 and c. 1000 respectively today. The Magnificent Frigatebird Fregata 
magnificens was still widespread in the islands during the late 19th century but today is 
restricted to two small islets, and only about five pairs are left. The traditional annual 
harvest of young Cape Verde Shearwaters Calonectris edwardsi has brought the total 
population down to c. 10,000 pairs (and probably less); it is estimated that 4-6000 young 
are taken every year. Cape Verde Petrels Pterodroma feae are taken locally because of the 
alleged medicinal properties of their fat. Apart from the Little Shearwater Puffinus boyd, 
the smaller procellariids are not actively persecuted but at times their colonies suffer 
greatly from the trampling of nesting burrows by fishermen who search the islets’ shores 
for shell-fish and debris. 

Due to the efforts of the NPPAP, some of the important sites for breeding seabirds 
were declared nature reserves by law in 1990. These include the islets of Raso, Branco, 
Ilhéus do Rombo, Curral Velho and Baluarte. Access to and utilization of these reserves 
are now subject to governmental authorisation, but law enforcement remains 
problematical on these remote and uninhabited islets. During the last decade, much effort 
has been directed at the implementation of educational and legislative programmes, 
aimed at the preservation of the remaining natural heritage of the Cape Verde Islands. 
But there is still a long way to go before an effective level of awareness of conservation 
issues is achieved, both among the general public and the responsible authorities of the 
country. 


The eight hundred and sixty-eighth meeting of the Club was held on Tuesday, 20 May 
1997 at 6.30 p.m., in conclusion of the AGM, in the Ante Room at Imperial College. 31 
Members and 8 guests attended. 

Members present were: The Rev. T. W. GLapwIN (Chairman), Miss H. Baker, 
Captain Sir THomas BarLow Bt. RN, I. R. BisHop, Mrs D. M. Brap ey, D. R. CALDER, 
Cdr M. B. CasEMENT RN, Professor R. J. CHANDLER, Dr R. A. CHEKE, D. J. FISHER, 
Professor C. H. Fry, F. M. GauNTLETT, D. GriFFIN, S. Howe, J. A. JoBLinc, R. M. 
Kett.e, Dr C. F. Mann, Dr J. F. Monk, D. J. Montier, Mrs A. M. Moors, Dr G. J. 
Moret, Dr M.-Y. More., Dr W. G. Portrous, Dr R. P. Pr¥s-Jongs, N. J. REDMAN, 
P. J. SELLAR, R. E. SHARLAND, Dr D. W. Snow, N. H. F. Stone, C. W. R. Storey, 
Dr R. WILKINSON. 

Guests attending were: Miss G. BoNHaM, Miss J. Cooper, S. DupDLEy, Mrs J. M. 
GLADWIN, Mrs S. GriFFIN, Mrs M. Montier, P. J. Moore, M. Paine, Mrs B. SNow, 
S. TONGE. 

After dinner, Members gave a series of short talks on subjects of topical interest. The 
following is a brief synopsis of the subjects discussed. 


Professor Hilary Fry gave a short talk entitled ‘““The amazing Anthoscopus nest’’. 
With the aid of a large sketch drawing, he described the unusual nest mechanics of the 
Penduline Tit. As co-author of The Birds of Africa (Academic Press), he has necessarily 
become involved in researching the literature ancient and modern, studying at the 
museum, and obtaining field data from colleagues. Integrating information from 
publications scattered in place and time enables him to give African birds a persona, 
often, it seems, for the first time. 

Suspended kapok nests of Africa’s Anthoscopus penduline tits, with their cryptic 
entrance slit at the end of the projecting spout, and snake-confounding false entrance, are 


Meetings 158 Bull. B.O.C. 1997 117(3) 


one of the wonders of the bird world (P. le S. Milstein 1975, Bokmakierie 27: 8-9). How 
the parent bird effects an entry has been described by several ornithologists—always 
rather differently. However, detailed observations by C. J. Skead in 1959 (Ostrich, 
Suppl. 3: 274-288) have now been verified by photographs, mainly by P. Steyn (1996, 
Nesting Birds, The Breeding Habits of South African Birds, Viaeberg). 

A parent arriving at the nest may mislead any watching predator by half entering 
the false ‘entrance’ (D. H. Chadwick 1983, National Geographic 163, 3: 344-385). Then 
it stands on the thick rim of the false entrance and, using its bill and one foot, pulls 
down the floor of the ‘real’ entrance spout overhead; it then levers itself inside, 
turns around in the spout, and draws together the spout ceiling and floor with its 
bill, whilst scrambling backwards into the brood chamber (Skead 1959). The coarse 
cobweb lining of the spout seems to promote adhesion. On leaving, the bird pushes its 
way along the spout, swings down, perches on the false entrance rim and with rapid 
movements jabs the underside of the spout upwards, with forehead and opened bill. It 
prods all of the underside, and then sometimes jabs also at the walls and floor of the false 
entrance. 

But this remarkable procedure prompts a number of questions: Does the false entrance 
concavity, with its greatly thickened ‘stoep’, also help to deform the spout entrance to a 
slit? Do structural stresses make the spout snap open or collapse, according to the bird’s 
position on, or in, the nest? And, finally, the kapok nest fabric is not woven, but is made 
into an extremély durable felt, by the tit’s jabbing and tweaking—how precisely is this 
achieved? 


Professor Richard Chandler gave a brief illustrated talk entitled “Sooty 
Oystercatcher Haematopus fuliginosus: two forms or two species?” 

Sooty Oystercatcher, an Australian endemic wader, like the other species of ‘black’ 
oystercatcher, is a bird of rocky shorelines. There are two forms, usually regraded as 
races, the nominate race occurring around much of western, southern and eastern 
Australia, while H.f. ophthalmicus occurs from Western Australia to Cape York. The 
status of the two forms is not clear. Hayman et al. (1986, Shorebirds: an identification 
guide to the waders of the world) suggest that ‘““Differences are marginal . . . the species is 
probably best considered monotypic. Most individuals cannot be assigned to either race, 
- and intergrading undoubtedly occurs.’’ In contrast, del Hoyo et al. (1996, Handbook of 
the Birds of the World) suggest that there is no field evidence for hybridisation between 
the two; “‘... the marked differences, especially in the bare parts, suggest that they may 
well prove to be separate species.’’ Recent field observations, in Victoria (Hf. fuliginosus) 
and at Darwin, N.T. and Broome, W. Australia (Hf. ophthalmicus) seem to support the 
latter view, and are presented here. 

The literature separates the two forms particularly by the broader, rather fleshy 
orange-yellow (rather than yellow-orange) orbital ring and the broader bill of 
ophthalmicus. There are conflicting views on which of the two has the longer bill. The bill 
breadth is not a good field mark, nor is bill length which seems to vary between 
individuals in both forms, presumably as a result of sexual dimorphism. There are 
differences in bill shape, typically uniformly tapered in fuliginosus, but in ophthalmicus 
usually rather deeper, particularly at mid length due to the increased depth of the lower 
mandible, giving an impression of an upturn in the bill. However, there is some overlap 
in shape between the two forms. These difference can be seen in the accompanying 
photographs, and can also be seen in photographs in Pringle (1987, The Shorebirds of 
Australia). 

Though the speaker admitted his experience is limited, these differences seem to be 
consistent. More fieldwork is clearly required, particularly to confirm (or otherwise) 
interbreeding between the forms. The differences are, in his view, ‘sufficiently great to 
support the view that they may be two species; at the very least, given reasonable views, 
they are easily separable in the field. 


Dr Robert Prys-Jones gave a short illustrated talk on ‘“‘Richard Meinertzhagen and 
fraud.”’ Following close examination of Meinertzhagen’s redpoll Carduelis spp. 
specimens, Knox (1993, Ibis 135: 320-325) made explicit the long-standing anecdotal 
rumours that this renowned ornithologist had fraudulently acquired and relabelled at 
least part of his large bird skin collection, most of which is now held at The Natural 
History Museum, Tring. Follow-up studies, including extensive, independent, 
radiographic evidence of the internal structure and make-up of skins both have largely 


Plate 2. (a) Sooty Oystercatcher Haematopus fuliginosus fuliginosus; Victoria, Australia, 
1 September 1996. Photograph: R. J. Chandler. (b) Sooty Oystercatchers Haematopus 
fuliginosus ophthalmicus; Northern Territory, Australia, 19 September 1996. Photograph: 
R. J. Chandler. (c) Hornby’s Storm-petrel Oceanodroma hornbyi; offshore southern Peru, 
27 October 1995. Photograph: W. G. Porteous. 


Meetings 159 Bull. B.O.C. 1997 117(3) 


confirmed the results. of Knox, and have extended his findings to encompass a diverse 
array of other species. As a result, locality and date of collection data attached to 
Meinertzhagen specimens should be treated with extreme caution. 


Joanne Cooper (of the Department of Geology, Royal Holloway, University of 
London.and Bird Group, The Natural History Museum) presented a short paper on 
“The Great Auk in late Pleistocene Gibraltar’. Remains of the extinct Great Auk 
Pinguinus impennis are known in small numbers from several late Pleistocene 
archaeological sites on the Portuguese and north Mediterranean coasts (Mourer-Chauviré 
& Antunes 1991, Geobios 2: 201-205). The presence of the species in these sites has been 
interpreted as evidence of hunting by man on its breeding grounds, thus indicating its 
summer breeding range during the last glacial period. However, recent investigations of 
the late Pleistocene avifauna from Gibraltar have suggested a different scenario for the 
Great Auk’s occurrence. Known from a total of five bones at three sites, it appears that 
the bones are more consistent with a pattern of scavenging of strandline carcasses by man 
or other mammalian predators, rather than hunting. It seems that this interpretation may 
be extended to other Mediterranean localities. 

Hence, its remains at these southern sites cannot be used as evidence of its late 
Pleistocene breeding distribution. Indeed, the exact opposite may be true, with these 
finds indicating a wintering distribution for the species. 


Dr Roger Wilkinson gave an illustrated talk on Vasa parrots. These are found only on 
Madagascar and nearby islands, and relatively little is known about their biology and 
behaviour. The genus Coracopsis comprises only two species: the Greater Vasa Parrot 
Coracopsis vasa and the Lesser Vasa Parrot Coracopsis nigra. Both are dull blackish or 
greyish brown. Vasa parrots are unique in showing seasonal changes in their bill, feather 
and skin colour, in everting their cloacas in the breeding season, and in their loud 
song-like calling. Studies of Vasa parrots at Chester Zoo have shown they have very short 
incubation periods, rapid chick development, and unusual sexual behaviour. 

The female Greater Vasa becomes bald-headed when breeding, and the bare skin on 
the top of her head, around her eyes and on her throat then turns mustard yellow. The 
copulation behaviour of Vasa parrots is unique among birds. Both sexes may show cloacal 
protrusions, which are most obvious in the male. The cloacal masses, when everted by 
the males, were found to be egg-shaped, 50-55 mm long xX 40-45 mm broad. The female 
less frequently everted her cloaca, which then appeared smaller than the male’s. During 
copulation, the cloacal mass of the male enters the female’s expanded cloaca, and the two 
birds remain locked together for lengthy periods—on one occasion this lasted over 100 
minutes! The Vasa parrot’s protracted cloacal contact is effectively a ‘‘tie’’ or “copulatory 
lock’’, similar to that which occurs in dogs. It seemed most likely that this is associated in 
some way with sperm competition, but the mating system of Vasa parrots in the wild 
remains largely unknown. Dr Wilkinson hoped that these studies may serve to stimulate 
field research in the wild. 


Dr Bill Porteous showed a series of slides, which he believed to be the first taken of a 
Hornby’s Storm-petrel Oceanodroma hornby1, photographed in the hand. This was one of 
the highlights of a voyage down the Humboldt Current, from Guayaquil, Ecuador, to 
Puerto Montt, Chile, in late October/early November 1995. The ornithologist on board 
for this tour was Peter Harrison, so the bird enthusiasts were in good hands. Hornby’s 
Storm-petrels were first seen near the Hormigas de Afuera islands, off the city of Lima at 
about latitude 12°S on 24 October, and were last seen off Arica in northern Chile at about 
18°S on the 27th. One individual came aboard the ship on the evening of the 26th and was 
closely examined, before being released the next morning. No observations were made 
between 18° and 24°S, due to land trips by day and night sailings, so the bird may occur 
further south than Arica, in October. 

From the literature sources he had consulted, it was clear that almost nothing is known 
about Hornby’s Storm-petrel. Its congener Markham’s Storm-petrel O. markhami is 
known to nest in southern Peru, but no active nest of O. hornbyi has ever been found. 
There is a report (but he had not been able to trace the original source) of mummified 
nestlings being found in holes in the northern Chilean desert at 1,500 m, 50 km from the 
coast. Hornby’s Storm-petrel occurs occasionally in concentrations of some thousands in 
the Humboldt Current off southern Peru and northern Chile, but since its breeding 
grounds are unknown it is impossible to estimate its population size or trends, or to define 


Meetings 160 Bull. B.O.C. 199% A473) 


conservation requirements. Further information about this elusive species would be 
welcome, from any source. 


Martin Gauntlett showed a slide of a Wandering Albatross Diomedea exulans, in 
flight, taken aboard the ex-Soviet Union research vessel MV Professor Multanovskiy 
(Marine Explorer), now being operated by ‘“‘Marine Expeditions’’ for Antarctic cruises, 
on a two-week trip to South Georgia and Antarctica, in 1995. It had long been his 
hope to see this species, and many were sighted on this trip, also numerous 
Black-browed, Royal, Grey-headed and Light-mantled Sooty Albatrosses. Since none of 
these kept a straight course for more than a few seconds, it was difficult to see how, in a 
recent paper in [bis (139: 234-252) which purported to show a correlation between flight 
performance of seabirds and wind direction, this could be calculated to any degree of 
accuracy, from measurements of their course and speed. One of the highlights of the trip 
was the landing on Albatross Island, South Georgia, having dodged the belligerent 
fur-seals to get ashore. There was an albatross nesting colony, with adults at the nest and 
almost fledged young from the previous year. To the surprise of the non-ornithological 
expedition leader, what the birders really wanted to see was the endemic South Georgia 
Pipit Anthus antarcticus. 


David Fisher decided that the next best thing to exhibiting a specimen of a new 
species, as used to be a tradition of the Club, was to exhibit what he believed to be the 
first and only sound recording of a species. He played a recording that he had made in 
Ecuador, in 1991, and asked whether anyone in the audience could recognize it. To 
everyone’s amazement, especially to the speaker, Bill Porteous immediately identified it 
correctly, as a Brown Wood-Rail Aramides wolfi. It transpired that Bill is one of the very 
few people who have seen this species in the wild, in recent years, and has published a 
note on his findings in Cotinga. David’s recording is to appear on a tape of New World 
rails currently being compiled by Bill Hardy and Bob Ridgely. 


Pat Sellar played recordings to illustrate the difference in song between Chiffchaff 
Phylloscopus collybita in England and P. canariensis in the Canary Islands. He used these 
examples to highlight the forthcoming BOU meeting on 3 December, when Dr J. 
Martens would explore the bioacoustic evidence supporting genetic differentiation in the 
Palaearctic Chiffchaff complex. 


Dr David Snow spoke about the winter song of Song Thrushes Turdus philomelos in 
his Buckinghamshire village, and the censusing of their breeding population over six 
years. In late October or early November a variable proportion of the males begin to sing. 
There is then a break in December, which in some years has coincided with the onset of 
a cold spell, but in other years has continued in spite of continuing mild weather. Song is 
resumed in early January by the same birds that sang in autumn, with additional birds 
beginning at any time from January to April. The final breeding population has been high 
(52-60 pairs in a village of a little over 100 acres), and suggests that the undoubted 
decline of Song Thrushes in farmland does not justify the alarm and concern for the 
survival of the species, that has been expressed in recent articles in the conservation 
literature and the national press. 


Ron Kettle played some tapes, and showed several slides of Hyacinthine Macaws 
Anodorhynchus hyacinthinus, taken on a visit to the Pantanal, Brazil, in August 1992. 
Groups of birds were photographed feeding together on their staple diet of palm nuts, 
and at nest sites often in association with nesting Jabiru Storks Fabiru mycteria. Sound 
recordings included their aggressive display calls, and from a group of 30-40 birds 
coming to roost in trees at dusk. 


David Griffin showed a series of slides of Firecrests Regulus ignicapillus caught and 
ringed in west London, and in particular a female with unusual plumage at Bedfont, near 
Heathrow, on 13 May 1995. A head-on view of this bird showed a well-defined, 
square-shaped black rib, bordered with white. No species of Regulus shows this feature, 
and the bird is therefore something of a mystery. Some have dismissed the bib as a patch 
of missing feathers, although the bib is very regular in shape. Others have suggested that 
this bird could be a hybrid—perhaps with one of the tit family? David Griffin invited 
comments. 

(Post-meeting note: John Marchant has examined these photographs and, in 
conjunction with the recorded wing and weight measurements, has concluded that it is a 
normal Firecrest, with a patch of feathers missing from its chin.) 


Accounts 161 Bull. B.O.C. 1997 117(3) 


British Ornithologists’ Club 
Financial statements for the year ended 31 December 1996 


Statement of Committee Members’ Responsibilities 


The Committee is required to prepare financial statements for each 
financial year which give a true and fair view of the state of affairs of the 
Charity at the end of the financial year and of the surplus or deficit for 
that period. 

In preparing the financial statements, suitable accounting policies 
have been adopted and consistently applied to the best of the 
Committee’s knowledge and belief, by reference to reasonable and 
prudent judgements and estimates. Applicable accountancy standards 
have been followed. 

The Committee members are responsible for maintaining adequate 
accounting records for safeguarding the assets of the Charity and for 
preventing and detecting fraud and other irregularities. ‘The 
Committee members are also required to indicate where the financial 
statements are prepared other than on the basis that the Charity is a 
going concern. 


Balance Sheet as at 31 December 1996 
1996 1995 


UNRESTRICTED FUNDS 

General Fund 
S2SCEE Dp TRAE" A RSE Snare ots ee mee ee eee eee 39,903 34,354 
SwiREUS Of LACOME OVET » PEXPENGITUTE ......<........00ccocsecscossnccsadeceseecosss 5,959 5,549 


Balance at 31 December 1996 45,862 39,903 


Herbert Stevens Trust Fund 
eta AE TP ATAUTAE LAO) <5. 20: a2 eee c Sade dat <donnwueccnsacos cccwcesaccsseuesncso8 OEM 162,791 143,841 
Net Loss (1995: Profit) on saies of investments during the Year......... (1,102) 8,629 
Increase in revaluation of investments at 31 December 1996 .............. 10,886 10,321 


Balance at 31 December 1996 12.575 162,791 


Barrington Trust Fund 
Balance at 1 January and 31 December 1996 577 577 


RESTRICTED FUNDS 
secre AEMaN TERME AVERED (SCE SIGLE2)) once ook sooo ndons cae eancne oie eieaceconcceaecdenss 3,867 3,330 


Bird Atlas of Uganda Fund (see note 2)..................scsccccesseeeseeeeeeeceeeees 2,597 — 
£225,478 £206,601 


Represented by: 
Fixed Assets 
EPEC EITC RCTNE oe BEG ci 50. c0seestesvsesessucecacedee tbo censde¥ehesciaiacsasesasresss 50 60 


Current Assets 
OES MEE AAES ELISEO TAS 52,85 0 soe n tzce TOTTI ae eA eee Oe belts fea Senehe 100 100 
Sy ARMMMEME SRLERCINS COM oe eh sic Seisaisi na vssuszdee Sune reas a ue Sceb TE adaeE SO evowsn Ue eROSeeS 30 38 

Cash at Bank: 
ERNE TAINS A CCOUNE sschciescs svaunuzcwsectbeeresctstaebex' tossiresésachoxcdeces 4,630 2,642 
—COIF Charities Deposit Account No. 1 .............cceccoccsssessssssseveees 50,324 45,576 
SLL ELLA C7 eee 2 See 2 ee in oP a ik) ee 141 782 


55,225 49,138 


Accounts 162 Bull. B.O.C. 1997 117(3) 


Current Liabilities 


Subscriptions received in advance ie/)).42.... sues cnecetenceaeo es cneee sh oducceused. ae. (4,740) (4,184) 
SUM GreditOrs cs. cceees Meare Mates sede at Bide cas owlee gen couse stow se ce ueu gon seawall (4,673) (5,111) 


45,812 39,843 


Herbert Stevens Trust Fund 
Investments at Marketi Value a. RE A aon 172,575 162,791 


Barrington Trust Fund 


111.57 COIF Charities Investment Fund Income Shares— 
at cost (Market value £871: 1995 £810) .0..........ccccceecccsececceeeeeeeeeeeees 577 577 


Publications Fund 
COIF Charities Deposit Account No 2 ...0........c:eccceeececneececneeeenseeseneees 3,867 3,330 


Bird Atlas of Uganda Fund 
COIF Charities Deposit Account No 3 .............cccceseeccceseeeaeeeceneeeeeeeees DOOM 


£225,478 £206,601 
Income and Expenditure Account for the year ended 31 December 1996 
1996 1995 
£ £ £ £ 
INCOME 
Subscriptions 
INTemibensi ere orca rte eeree rere oe ee ee OTe ere RR nc Ha Hau aI BE CLE 4,961 4,893 
Non-member, subscribers ....00.6 oosccenctlinnslgje soduoes  Saccedudea tiles teeuaceaqdehcsemeardes 3,313 2,762 
Income Tax recovered under Deeds of Covenant .................000c00ec00e00= 308 311 
8,582 7,966 
Donations 48 20 


Investment income 


Herbert Stevens; Brust Butrico oss sito ae eokuesatciceaaaem sae secon asmcadek dose aueams 9,912 7,090 
Barrington Trust Fund (COIF income Shares) ..................ccesseeeeeeeeees 38 36 
Interest Received: 
Barclays Pritme Account. 63.55. eba et ieecss sens sone nuecou nobetduie » dedeeteaencueeds t 139 125 
COEF Deposit Account INO ier tek ce cones sce eueneeseneccusscuesceecanen sees 2,698 2,561 


12,787 9,812 


Sales of Publications 
BOC Bulletin: 


IBAGKINUIM DE TS shook eancaeeaatnuacarncesetoenee ae ee aaeakiee aineaitealglauual satalauemeattae 1,858 469 
OP rit york. seaetan steko oeseaen sera Pesaemeen oc eae doko sae ameneenaet scam 96 226 
Contributions) to! Colour: Blates)..): acct sweeecesecass-+ oecceseee = cpaene taco —_ 584 
TMSENiS% see tances nnwnincoatemt: atucs 2 dhl Meas Wena Sear Gees cam con nh senna aaa 160 55 
“Avian Systematics and Taxonomy ’’...............ccssseeeeccneseeeeensneceeenees 849 TN itS)S) 
“Birds, Discovery and Conservation”? .............cc0cceseecneccnseeeseeneeeasenes 236 134 
SExtinct/and! Pndangered Birds’ > 4-20. 2c2csccc--coceskete-cersesestereee sees ease 373 1,458 


3,572 4,081 


Cost of sales 


OpeningaStockiise thie. Me teets 5 eM ches demaset aes sce ten s Maaants satel a (100) (100) 

Closing Stocks. 09, sdtncgs2on«ctedeies satraauidnap ti evasiasaets ogee yltead detteue shia asladece stearate 100 100 
Meetings 

Ordinary Meetings 2.2 ysl. Mates Site newawdugavanendousendamaeddeayseranveeands 3,131 3,470 

CentenarysDintieniyt 2.5 )t hte s. ss eis SO sa OR Ry eae _ 8 


3,131 3,478 


Accounts 163 


Miscellaneous income 


CIPS Oa ol 2 OS, ee ee eee eee cee 
Set ORS ey a ene Rea et RT RE ES Sak Oe 


TOTAL INCOME 


EXPENDITURE 


SNR ERE ERYS ENON IMEI UCASE UNE ESTA © ERE Gof c321 5 50555 Gace de «sis os oa ste ok Sac cnekesedeseeesicaccee 


Meetings 


ER CEIMMMET STEP ROO TIL EDIE E esi sass oa cc coc aedde as cosas Shas agnaeeatesoaagaasadeanensddvecasone 
irepestahitiy CHAT DOS 2556 So es EE soc cen atc souvaauasexsouns Meoseressaestes 
STRIAECS INIATCES Breck 2 SEE nee cere Renee es ener ene eee 


BOC Bulletin 


Pama NSEC eA TOA SAENGS) PNT AUD UATD 03 2 655 ve 5s coe anos nds svn cea ecdeweceu sex ebecesiuccsyeceses 
Additional offprints and back numbers....................ceeeeeeeeeeeeeeeeeees 


LE ES PESTS POET EEE 1g ee er 
Banorial and secretarial EXPENSES 65.2. .<ss..00.s22sssdesevessncoveusescccsssseds 
WARMERS SPE AENE ES Ce ooS oct n coco ct ox conn vs sass tosses ccoaussnacdess sedesusdeseecseessseteade 


Other publications 


“Birds, Discovery and Conservation” purchased .................0e0ee000 


“Extinct and Endangered Birds” 


LESEDTEE, Preeseeti £ [AU ES DL OE or a eT 


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Publications Sub-Committee expenses .................cseeeeeceeeeeeeceeeeees 


DEPRECIATION (Projection Equipment) ....................ccseccesesseeeees 


Miscellaneous 


Rg NRINIE CRE AIAEAT ERAS UTA 8 3 os oo a etiay can ps duasoh snanssoanesBeeos jay eve sa eaheo 
PECOMEANCY TEES: OFUIATY TUNGS -.......0.<cc0pccnsosssescacssesesveccoussnvessasees 
RaaG HEAT: WET —DTOVISION .......0.0cs50ccecnastrestectscerecstocesce otenee tee 
Rickie pes Mrtist Pini |: 522520223 0cc02 9.0400 8evoeden sp guveveaessbuccezs 
Legal advice: Herbert Stevens Trust Fund...............ccccccceeeeceeceeedes 
Bank Cunnpes: font! .lepetion) 25) ioe. Stes 2oos: 
Credit card chargespecsc.cezs5. 10-0! 2. dace ugyed paris oo. pesdadn eceeso nsec? 
Binding Club copy of Bulletin back numbers ................2s0see0eeeeee00s 
ELD ET 27 TEE Ae SE RR ROE, Se Pine eee ee 


CLL SEPT SLL ye ee ceeatene tial oe iinet il uch axl Mion nll ate ae 


21 


13 


418 


216 


Bull. B.O.C. 1997 117(3) 


90 


28,210 


1996 


3,454 


13,178 


897 


1,555 
277 


10 


133 


2,805 


£22,251 


67 
167 
40 


492 


146 


274 
£25,631 


1995 


3,922 


11,633 


1,109 


1,621 


133 


1,506 
£20,082 


Accounts 164 Bull. B.O.C. 1997 117(3) 


Excess of Income Oven EXPeNditune cy creecesscce-cssecesee ons ome eace teste seeeee eens £5,959 £5,549 


NOTES TO THE ACCOUNTS 
1. ACCOUNTING POLICIES 
(1) Basis of Accounts 
The financial statements are prepared under the historical cost convention. 
(2) Depreciation 
Depreciation is calculated to write off fixed assets over their expected useful lives, by reference to original cost or 
subsequent valuation at the following annual rate: Projection Equipment, 10% on cost. 
(3) Stock 
Stock is valued at the lower of cost and estimated net realisable value. 


Bird Atlas of 


2. RESTRICTED FUNDS Publications Fund Uganda Fund 
1996 1995 1996 
Balance ath Namuary 1996 uo caste ee net acne ces sneer cece oar neeoeec mecca eee 3,330 —_— — 
Sponsorship received iy sis cqakecc sees. Ab ceachanc. v0 oosdvecataawaes sevenvnnvedescseceataeseess —_— — 2,488 
DJ ONAtTOTIS Sra Sere eles Ramet sates ee ee ah ca cs cted tase eeateaeceoe smc esaeeemeaeteees 337 2,400 —_ 
Income: TaxtRecovere dic. .cs.c:.tsncdee Peeee cos. sons Rena eecsertwadaayseacasenevsetaddtarsees —_ 800 —_— 
Interest Gross enc ce sree tata erect core eee ee oer aRs aaNd eee Shes ae ame eR RO oe 200 130 109 
Balance at 31 December 1996 £3,867 £3,330 £2,597 


HERBERT STEVENS TRUST FUND 
Balance Sheet as at 31 December 1996 


1996 1995 
£ £ 
General Fund 5: 
Balancetatiaianuany MiG 96 Rs ice snvearsin. oh cdeoeataneaim evan cheancoumieaaereccdetec es 162,791 143,841 
Loss (1995: Profit) on sale of investments during the year....................- (1,102) 8,629 
Increase on Revaluation of Investments at 31 December 1996............... 10,886 10,321 
Balance at'3il "December N99 Ger. tse ees see rc cenes ov ao neee mectnecoesccemeeeeens £172,575 £162,791 
Represented by: 
Investmentsatmarket value. .....0:00:...c.<.c2cssescssscssuseeesaedaeaecsdesucsseedeoss< 1729575 131,855 
Current Assets: Midland Bank SARA ..................ccccscceccneeeneesecceneceees — 30,936 
£172,575 162,791 


INDEPENDENT EXAMINER’S REPORT 
TO THE COMMITTEE MEMBERS OF 
THE BRITISH ORNITHOLOGISTS’ CLUB 


We report on the accounts of the Charity for the year ended 31 December 1996. 


RESPECTIVE RESPONSIBILITIES OF TRUSTEES AND EXAMINER 

As the charity’s trustees you are responsible for the preparation of the accounts; you 
consider that the audit requirement of section 43(2) of the Charities Act 1993 (the Act) 
does not apply. It is our responsibility to state, on the basis of procedures specified in the 
General Directions given by the Charity Commissioners under section 43(7)(b) of the 
Act, whether particular matters have come to our attention. 


BASIS OF INDEPENDENT EXAMINER’S REPORT 

Our examination was carried out in accordance with the General Directions given by the 
Charity Commissioners. An examination includes a review of the accounting records kept 
by the charity and a comparison of the accounts presented with those records. It also 
includes consideration of any unusual items or disclosures in the accounts and seeking 
explanations from you as trustees concerning any such matters. The procedures 
undertaken do not provide all the evidence that would be required in an audit, and 
consequently we do not express an audit opinion on the view given by the accounts. 


Accounts 165 Bull. B.O.C. 1997 117(3) 
~ INDEPENDENT EXAMINER’S STATEMENT 


In connection with our examination, no matter has come to our attention: 

(1) which gives us reasonable cause to believe that in any material respect the 
requirements 
—to keep accounting records in accordance with section 41 of the Act; and 
—to prepare accounts which accord with the accounting records and to comply 
with the accounting requirements of the Act 
have not been met; or 

(2) to which, in our opinion, attention should be drawn in order to enable a proper 
understanding of the accounts to be reached. 


Prince Albert House 
20 King Street DONALD REID & CO., 
Maidenhead, Berks Chartered Accountants 


30 April 1997 


Approved by the Committee on 20 May 1997 
D. GRIFFIN, Chairman 


A. Madrotio N. & E. Z. Esquivel 166 Ball. B.O.C. 199 7enG) 


Noteworthy records and range extensions of 
some birds from the Reserva Natural del 
Bosque Mbaracayu (Mbaracayt Forest Nature 
Reserve), Departamento de Canindeyu, 
Paraguay 


by Alberto Madrono N. & Estela Z. Esquivel 
Received 12 October 1995 


Introduction 


The Reserva Natural del Bosque Mbaracaya (RNBM or Mbaracayt 
Forest Nature Reserve) is one of the few remaining areas of humid 
subtropical Atlantic Forest in Paraguay. This “‘Atlantic-type”’ forest is 
a westward extension of the true Atlantic forest of southeastern coastal 
Brazil and is sometimes referred to as “‘Bosque Paranaense’’ (Cabrera & 
Willink 1973, Laclau 1994), “‘Bosque Atlantico del Interior’ (FMB 
1994) or “Brazilian Interior Atlantic forests’’ (Dinerstein et al. 1995). 
We refer to it as the Paraguayan Atlantic forest for being part of the 
Endemic Bird Area of the same name (EBA B-52: see Wege & Long 
1995, also ICBP 1992). The remaining Paraguayan Atlantic forest has a 
tremendous conservation value as it still holds populations of several 
threatened and near-threatened birds, including some endemic to the 
Atlantic forest (Brooks et al. 1993, Madrono N. & Esquivel 1995, 
Lowen et al. 1995, 1996a,b). 

The RNBM is centred on 24°08’S, 55°25’W in the Departamento de 
Canindeyt, northeastern Paraguay, c. 10km east of the town Villa 
Ygatimi. It has an almost rectangular shape of 57,715 ha with the 
northeastern corner contiguous to the border of Mato Grosso do Sul, 
Brazil. On the eastern side, another adjoining fraction of 5487 ha of 
mainly campos cerrados brings the total protected surface to 63,202 ha. 
The greater part of the reserve (c. 85%) is forested (tall, medium, low, 
flooded and gallery forests), although there are other natural 
communities as well, including grasslands, marshy areas, lagoons etc. 
To date, no detailed botanical inventory work in the reserve has been 
compiled other than CDC’s (1991) general description of its different 
natural communities and Keel et al. (1993) who studied tree and shrub 
composition of several sample plots at one locality within the RNBM, 
concluding that the area studied was a top conservation priority in 
eastern Paraguay. 

Typical trees of the higher canopy include: Astronium fraxinifolium, 
Aspidosperma polyneuron, Tabebuia heptaphylla, Albizia hasslert, 
Peltophorum dubium, Anadenanthera colubrina and Balfourodendron 
riedelianum. Some of these trees can reach heights of up to 30-35 m. In 
a medium stratum the tall forest has trees up to 20-—25m such as 
Chrysophyllum gonocarpum, Diatenopteryx sorbifolia, Helietta apiculata, 
Myrciaria sp., Holocalyx balansae, several species of Lauraceae, 


A. Madrotio N. & E. Z. Esquivel 167 Bull. B.O.C. 1997 117(3) 


Cabralea canjerana and introduced Citrus aurantium. 'The low strata are 
mainly composed of shrubs that normally reach 4-8 m. Common 
species at this level are Sorocea bonplandiu, Inga marginata, Pilocarpus 
pennatifolius, Trichilia spp. and Piper spp. The ground cover varies 
considerably from one place to another, but ferns of many types, 
Rubiaceae, Bromeliaceae and grasses are very common (more details on 
different habitat types in the reserve will be published in Lowen et_al. 
in prep.). In the main text, we also often refer to cerrado of Aguara Nu, 
this being characterised by a mosaic of savanna (with varying 
concentrations of Butia palm), pure grasslands, and to a lesser extent 
xerophytic woodlands that generally surround the more open areas. 

The RNBM is drained to the west by the upper Jejui’mi river, a 
tributary of the Paraguay river. It is an area characterised by gently 
rolling hills mainly composed of soft sedimentary rock weathered into 
sandy soils of low fertility. There are also (in less extent) patches of red 
clayish soils which are generally more fertile and more commonly 
found to the east in the Parana river basin. The entire area is generally 
between 150-300 m above sea level. The annual rainfall is 1600- 
1800 mm (1800 mm in 1995 at Jejui’mi), with a dry cold season 
(between July and September) and a wet hot season (between October 
and February). T‘emperatures are highest during the summer months 
(December—February), with an average daily temperature of 27°C in 
January and maximum temperatures unlikely to surpass 38°C. During 
typical winter months (July and August) daily temperatures average 
17°C, but temperatures can drop considerably (even below 0°C, with an 
average of 4-5 frosts each year; Acevedo et al. 1990). 

Most of the information provided in this article is the result of 162 
days of fieldwork completed between July 1994 and June 1995, which 
in most cases was conducted by both authors together (c. 300 man-days 
of field work and c. 1800 man-hours in the field). Fieldwork was 
distributed evenly over the year in order better to understand 
year-round changes in the bird community composition and activity 
patterns (to be published). Most bird species breed mainly in spring 
(September—November), but our knowledge of breeding seasons of 
many species in the reserve is still very limited. Records provided by 
other ornithologists are fully credited in the text. For a few species we 
cite records obtained after June 1995. For those species on which we 
give our opinion regarding their status in the RNBM (e.g. uncommon, 
rare, etc.), this is based upon our experience in the field up to February 
1996 (which in total includes some 200 days in the field for each of 
the authors. 

This paper summarises some of the most interesting records during 
this period. They have been chosen for their importance in 
contributing to current knowledge of the species’ distribution or status, 
or because there are few records either in the entire country, or in 
Oriental Paraguay (““Orient’? in Hayes 1995), ‘“‘Central Paraguay”’ 
geographical region (as defined by Hayes 1995: 19) or the Dpto. 
Canindeyu. Some of the species are threatened or near-threatened 
(following Collar et al. 1994), in which case T (threatened) or NT 
(near-threatened) has been added in brackets after the scientific name. 


A. Madrotio N. & E. Z. Esquivel 168 Bull. B.O.C. 1997 117(3) 


Although not necessarily mentioned under each species account, our 
knowledge of previous records is mostly derived from the exhaustive 
monograph on the birds of Paraguay by Hayes (1995), whose 
taxonomic order is followed. Readers are also referred to Madrono N. 
& Esquivel (1995) who provide additional noteworthy records for the 
RNBM that complement the information presented here. 

Coordinates (latitude and longitude) are given for each record, and 
have been obtained either using a Geographic Position System 
(Pathfinder Basic Plus, ‘Trimble Navigation) or from a 1:50,000 scale 
map (Mapa Geografico Militar) in combination with black and white 
1:50,000 aerial photographs (see below). Coordinates without seconds 
(") have been obtained from the map and we are certain that the error 
should not exceed +500 m. For visual records, we used Zeiss 10 X 40 
and Vision 10 X 50 binoculars. Unless otherwise stated, records below 
were made by both authors. Place names are followed by coordinates 
the first time they appear, but not when mentioned later. 


Species accounts 
HOOK-BILLED KITE Chondrohierax uncinatus 


There are few records from Oriental Paraguay; the records below 
are the first for Dpto. Canindeyu. One bird seen by AMN in flight on 
16 September 1994 (24°07'54"S, 55°31'36”"W), and another (possibly 
the same) on 29 Sept 1994 (24°08'03"S, 55°31'44”W). Further obser- 
vations in August and September 1995 suggest that breeding may occur 
in RNBM. 


RUFOUS-THIGHED HAWK Accipiter erythronemius 

Neither Hayes et al. (1994) nor Chesser (1994) include this species as 
an ‘“‘austral migrant’’ (either “northern austral migrants’, 1.e. birds that 
breed in Paraguay and migrate north in winter, or “southern austral 
migrants’’, i.e. birds that nest farther south and winter in Paraguay). 
On 10 March 1995, no less than 15 birds were observed flying north 
(c. 24°07'S, 55°31’W), with an obvious migratory behaviour (AMWN and 
P. Donahue). Cold light winds from the south seemed to facilitate 
migration as other raptors, such as 12 Swallow-tailed Kites Elanoides 
forficatus, were also part of the same loose mixed group. 


BLACK-AND-WHITE HAWE-EAGLE Spizastur melanoleucus (NT) 

Hayes (1995) lists up to four records for Oriental Paraguay. The 
unspecified observations of the species in RNBM (Madrono N. & 
Esquivel 1995) are as follows: one bird soaring close above the forest 
canopy on 3 September 1994 at 24°07'29"S, 55°26'43”W (AMIN) and a 
perching bird on 26 May 1995 at 24°07'52"S, 55°31'26"W (EZE). 
Another perching bird was observed at Jejui’mi (24°08'03"S, 
55°31'44”W) on 6 April 1995 (P. Donahue). The species seems to be 


rare in the reserve, where the authors have only recorded it twice. 


ORNATE HAWKE-EAGLE Spizaetus ornatus 


There are many records for the country, but none made with 
certainty since 1939 (an undated record from Parque Nacional Cerro 


A. Madrofo N. & E. Z. Esquivel 169 Bull. B.O.C. 1997 117(3) 


Cora is the only possibility; see Hayes & Scharf 1995), and none has 
ever been recorded in Dpto. Canindeyt (see Hayes 1995). The species 
has now been recorded at RNBM, where a bird was observed flying 
high, calling, on 14 and 15 February 1995 at Horqueta’mi (24°08'10"S, 
55°19'21”W) by both authors and forest ranger S. Ramirez, and on 
11 April 1995 at 24°07'37"S, 55°31'01"W by AMN. Indigenous Aché 
hunters know the species from the area, reporting previously finding at 
least one nest. The scant number of records suggests that the species is 
rare in the reserve. 


BLACK HAWK-EAGLE Spizaetus tyrannus 


Hayes (1995) regards the species as “‘hypothetical’’ for the country, 
adding that ‘‘further documentation is needed before this species can be 
accepted for Paraguay’’. Our records at RNBM confirm its occurrence 
in Paraguay (unless otherwise stated, observations took place at Jejui’mi 
in 1995 and refer to a single bird calling in high flight): one silent bird 
carefully observed over five minutes on 25 October 1994 at 24°08’40"S, 
55°31'34"W (AMN, D. Pullan and R. Denny); two birds seen soaring 
high together in November 1994 (R. Clay); briefly observed on 17 
March; 20 May (AMN), 21 May, 3 June (AMN), 13 June (forest ranger 
N. Lopez) and 20 June (low flight, AMN) 1995. An additional record 
of the species occurred at Lagunita (24°08'04"S, 55°25’42”W) on 13 
September 1995, where a bird was observed and its call tape-recorded 
(AMN, EZE and many other ornithologists of the joint Anglo- 
Paraguayan “‘Project Yacutinga ’95’’). 

A local resident, E. Caballero (now working at RNBM for the 
Fundacion Moisés Bertoni), informed us that a few years ago, while 
sitting in a mangrullo (a hunting platform) south of Lagunita (c. 
24°09’S, 55°25'W), he observed a large black eagle (presumably a Black 
Hawk-Eagle) that took a Brown Capuchin monkey Cebus apella from 
the upper canopy. 

Birds have been heard giving a single long whistle lasting about 1.5—2 
seconds, rising slightly and then falling quickly at the end. Another 
variation incorporated three fast short whistles preceding the longer 
call described above or interspaced between two long calls, this 
resembling the description given in Sick (1993) and also birds heard in 
Brazil, Ecuador and Panama (R. Clay verbally). Additionally, full 
descriptions were taken and the birds conformed to the literature, 
notably fitting Canevari et al.’s (1991) colour illustration of the species. 


SUNGREBE Heliornis fulica 


There are only two confirmed records and five undated, unconfirmed 
reports in Paraguay, all from Oriental Paraguay. Acevedo et al. (1990) 
list the species as if it was present in RNBM, but this was probably a 
guess, the species not being listed for the reserve in any of the 
unpublished lists (e.g. CDC 1991, FMB 1992). We have, however, 
observed/heard the species on several occasions at the Jejui’mi river 
between 24°08’S, 55°31’W and 24°10’S, 55°30’W. On 23 November 
1994 an adult bird was persistently calling, and also observed, at 
24°09'33"S, 55°30'24”W. The species seems to be uncommon, although 


A. Madrotio N. & E. Z. Esquivel 170 Bull. B.O.C. 1997 117(3) 


probably has a healthy population along the Jejui’mi river within the 
reserve. 


RED-LEGGED SERIEMA Cariama cristata 


Hayes (1995) does not include the species for Central Paraguay. It is 
present, however, in the Aguara Nu cerrado in the easternmost part of 
the reserve (mainly 24°08-17'S, 55°15-17'W). Recently, after a large 
fire took place in the above-mentioned area, a single bird was observed 
on 12 and 14 October 1994 (S. Ramirez and J. Padwe verbally) on the 
main unpaved road that crosses the RNBM west to east, in an area 
surrounded by tall forest (at 24°07'S, 55°28’W), c. 20 km away from the 
species’ habitual grounds. The presence of the bird in such habitat 
(although admittedly on the road) is exceptional (the indigenous Aché 
had never seen this species in the forest) and presumably was due to the 
fire. 


SCALED PIGEON Columba speciosa 


Hayes (1995) lists only three records for Paraguay (all from Oriental 
Paraguay). Although the species was already recorded at RNBM 
(Brooks et al. 1993), we have now observed it year-round on many 
different occasions throughout the reserve, in forests and also islets of 
forest in the flooded grasslands of the Jejui’mi river. We judge the 
species to be uncommon but with a healthy population within the 
reserve. 


HYACINTH MACAW Anodorhynchus hyacinthinus (‘T) 


All bona fide records of this species in Oriental Paraguay are from 
Dpto. Concepcion (see Hayes 1995). A record of “‘some birds” south of 
Parque Nacional Cerro Cora (questioned by Lopez 1992 and hence by 
Hayes 1995) refers to captive birds of uncertain origin (F. Colman 
verbally 1995). The three records reported in Madrono N. & Esquivel 
(1995) extend the species’ known range about 200 km to the southeast 
into Dpto. Canindeyu. The most recent record at RNBM occurred in 
Aguara Nu, in an area of cerrado vegetation at 24°10'49"S, 55°15'34"W 
on 18 October 1994, where a single bird was observed by the forest 
ranger J. C. Almada. These records in Canindeyt suggest that the 
species perhaps undertakes occasional movements (away from the 
extensive cerrado of Concepcion) to other scattered and smaller cerrado 
existing further south (Dptos. of Amambay and Canindeyt) that 
provide seasonal fruiting of the jatai palm Butza sp. (although this fruit 
has never been reported as part of its diet: Collar et al. 1992, N. E. 
Lopez verbally 1995). That the species undertakes seasonal long- 
distance movements has already been mentioned (Collar et al. 1992), 
although little information is available. 


PHEASANT CUCKOO Dromoccocyx phasianellus 

The following records at the RNBM add Dpto. Canindeyt to the 
scant number of known localities for this species in Paraguay. A bird 
was heard (and tape-recorded) on 27 October 1994 at 24°07'20"S, 
55°31'41"W (AMN, D. Pullan and R. Denny); one was observed on 


A. Madrotio N. & E. Z. Esquivel 171 Bull. B.O.C. 1997 117(3) 


16 March 1995 at 24°08’S, 55°31’W; one was heard on 29 August 1995, 
1 and 11 September 1995 at 24°07'37"S, 55°31'01"W (AMN). The 
records above suggest that the species is a rare year-round resident in 
the area. 


MOTTLED OWL Ciccaba virgata 

There are only two recent records of this owl in Paraguay; Brooks 
et al. (1993) recorded it once at the RNBM (sight record). Since then, 
we have heard its call many times in high and medium forests in 
different parts of the reserve throughout the year. 


OCELLATED POORWILL Nyctiphrynus ocellatus 

Hayes (1995) only lists four records for Paraguay rata in 1978 from 
Dpto. Canindeyt; Storer 1989). We have now recorded the species 
many times throughout the year (1994—1996) in different areas of the 
RNBM (most are aural records). Breeding was also confirmed on 
27 October 1994 at 24°07'35"S, 55°31'45”W, when an incubating adult 
(with a single egg) was found on a path in low forest close to low 
flooded forest (R. Denny, D. Pullan and AMN). The species seems to 
be uncommon in the reserve (although we believe a healthy population 
exists), most of the encounters being in tall forest away from the path. 


SILKY-TAILED NIGHTJAR Caprimulgus sericocaudatus 


-'There seem to be only three published records for the country (see 
Storer 1989). We have heard the species on several occasions at the 
RNBM: 14 July 1994 (24°08'33"S, 55°20'26”"W); 17 and 18 August 
1994, 19 November 1994 and 14 February 1995, all at Jejui’mi; and 
16 March 1995 at 24°08’S, 55°31’W. The above records, plus another 
bird heard at Lagunita on 13 September 1995 by AMN, show that the 
species is an uncommon year-round resident in the RNBM. 


LONG-TAILED POTOO Nyctibius aethereus 

Although twice recorded at the RNBM (see Hayes 1995, Lowen 
et al. in prep.), there are still relatively few records for the country of 
this secretive species. On 11 February 1995 we had very close views of 
a silent bird perching and flying (the observation lasted more than 
15 minutes) in the cerrado area of Aguara Nu at 24°10'31"S, 
55°15’32”W. Between late August and November 1995, a pair was 
breeding at 24°07'38"S, 55°31'08”W (incubating adults, a nestling and 
fledged juvenile were observed). 


GREY-RUMPED SWIFT Chaetura cinereiventris 


Hayes (1995) indicates that the species is ‘“‘apparently present during 
winter’. We have observed Grey-rumped Swifts all year round at the 
RNBM. 


GREEN-AND-RUFOUS KINGFISHER Chloroceryle inda 


Hayes (1995) lists four records for Paraguay, the only recent records 
being from the RNBM in 1992 (Brooks et al. 1993). We have found 
the species regularly throughout the reserve year-round. Our records 


A. Madroto N. & E. Z. Esquivel 172 Bull. B.O.C. 1997 117(3) 


also include one active nest (with a minimum of two chicks) found on 
29 November 1994 at 24°12'23"S, 55°29'53”"W. 


PYGMY KINGFISHER Chloroceryle aenea 


There are only two previous records for the country (Hayes 1995). 
The finding of the species at the RNBM is the first record in Oriental 
Paraguay: one bird was briefly observed on 5 November 1994 (EZE), a 
pair on 11 November 1994 (R. Clay), and one bird on 7 April 1995 
(AMN, EZE, P. Donahue and T. Wood). All records were at c. 
24°08'S, 55°32’W. 


COLLARED CRESCENTCHEST Melanopareia torquata 


There is just one record for the country, from 1938 in Amambay 
Department (Ridgely & Tudor 1994). We observed one bird in short 
grassland in the cerrado of Aguara Nu on 9 February 1995 at 24°10'S, 
55°15’'W. Breeding in the area seems likely as further observations 
between 14 and 20 September 1995 by the authors and several other 
ornithologists of ‘“‘Project Yacutinga ’95’’ showed that several pairs 
were present (Lowen et al. in prep.). 


SHARP-TAILED TYRANT Culicivora caudacuta (NT) 


The species has not been recorded in Oriental Paraguay since 1932 
(Hayes 1995). We observed two birds (adult and immature) in 
grassland at the cerrado of Aguara Nu on 13 February 1995 at 
24°09'29"S, 55°17'21” W. Breeding in the area would seem likely, with 
further observations of the species by the authors and several ornithol- 
ogists of “‘Project Yacutinga ’95”’ (one bird mist-netted and photo- 
graphed) between 14 and 20 September 1995 (Lowen et al. in prep.). 


BAY-RINGED TYRANNULET Phylloscartes sylviolus (NT) 


Madrono N. & Esquivel (1995) reported not having found this 
species after 162 days of fieldwork. It has been previously recorded in 
the reserve (see FMB 1992, Brooks et al. 1993). AMN observed the 
species on 29 August 1995 at 24°07'37"S, 55°31'01”W, a site where a 
pair and single birds have subsequently been observed regularly. 
Earlier in August 1995, D. Finch (verbally 1995) discovered a pair 
building a nest somewhere to the east (not further than 1 km from the 
above coordinates). The species is uncommon in the reserve, but 
probably has a healthy population. It is certainly difficult to detect, due 
to its habits of moving about high in the canopy and its low-pitched 
call. It is noteworthy that many of AMN’s observations occurred 
in exactly the same place in a Copaifera langsdorfi tree between 
August 1995 and January 1996, suggesting some degree of seasonal 
territoriality. 


LARGE-HEADED FLATBILL Ramphotrigon megacephala 


There are just two recent records for the country, one of which was 
at the RNBM (Hayes 1995). We found the species twice in mixed giant 
bamboo Guadua sp. growth on 15 June 1995: one bird heard at 24°15'S, 
55°21’W and one seen and heard at 24°15'49"S, 55°22'04"W. The 


A. Madrono N. & E. Z. Esquivel 173 Ball? BOC. 1997 M17) 


species was again present at the first locality on 22 and 23 September 
1995 (Lowen et al. in prep.). 


CURL-CRESTED JAY Cyanocorax cristatellus 

In Paraguay, the species has only been reported in Dpto. Concepcion 
(Ridgely & Tudor 1989). It has now been recorded at the RNBM, with 
the first record in winter, probably August 1993 (forest rangers J. C. 
Almada, N. Lopez, T. Osuna, S. Ramirez and R. Villalba). After this 
sighting, the species has been observed repeatedly in the same area, 
with our first observation, between 9 and 12 February 1995, all in the 
cerrado area of Aguara Nu at different localities (same general area 
mentioned above under the Red-legged Seriema). Most of the records 
refer to small groups of up to ten birds. One exceptional record of two 
birds occurred probably in December 1994 at Jejui’mi, in a small 
clearing surrounded by tall forest c. 22 km west of the species’ habitual 
site at Aguara Nu (S. Ramirez). ‘Two Guarani indians inhabiting part 
of the Aguara Nu area independently informed us that the species has 
recently colonised this area from Brazil. It was previously only known 
to them from over the border in Brazil and thus they named it “‘Aka’é 
Brasil” (“‘Aka’é” is the generic Guarani name for the two other species 
of Cyanocorax present in the area). 


BANANAQUIT Coereba flaveola 


The species was not previously recorded from Dpto. Canindeyt. 
Records at the RNBM are as follows: one bird on 8 March 1995 
(P. Donahue), 13 and 14 March 1995 (AMN and EZE), all at rio 
Jejuir’ mi (24°08'32"S, 55°31'34"W),. 


SILVER-BEAKED TANAGER Ramphocelus carbo 

The species has only been reported twice in Oriental Paraguay 
(Hayes 1995). A bird was recorded on 10 March 1995 for the first time 
at the RNBM., at the rio Jejui’mi (P. Donahue, AMN and EZE). 


TEMMINCK’S SEEDEATER Sporophila falcirostris (T) 

This rare Atlantic Forest endemic bamboo specialist has only 
previously been recorded in Paraguay in July 1977, west of Saltos del 
Guaira, Dpto. Canindeyt (Ridgely & Tudor 1989). It has now been 
found in the southernmost area of the RNBM, where giant bamboo 
growth is the most extensive. On 15 and 16 June 1995 (note winter 
date), a singing male was observed at 24°15'52"S, 55°22'02"W, and a 
second singing bird was heard nearby at 24°15’50"S, 55°22’23"W. A 
third singing bird was heard more than 1 km away (to the west). All 
birds exclusively used giant bamboo (Guadua sp. not “‘Bambusa sp.’’ as 
erroneously stated in Madrono N. & Esquivel 1995), and a male was 
observed eating what were presumably bases of bamboo leaf petioles. 
Bamboo was certainly neither flowering nor seeding. Only these three 
birds were detected despite our walking a minimum of 5 km through 
habitat apparently suitable for the species. Further searches at the 
above-mentioned localities on 22—23 September 1995 during ‘‘Project 
Yacutinga ’95”’ and in October 1996 did not locate the species. 


A. Madrotio N. & E. Z. Esquivel 174 Bull. B.O.C. 1997 117(3) 


The survival of the species in the area is in doubt, considering its 
strong association with giant bamboo forests. Suitable habitat for 
‘Temminck’s Seedeater in the reserve is relatively restricted (perhaps no 
more than 5000 ha). The protection of an adjoining fraction of this type 
of mixed-bamboo forest south of the reserve is judged crucial to the 
species’ long-term survival. This land is certainly the last tract of 
undisturbed forest around the reserve, with perhaps some 10,000 ha 
(privately owned). Other threatened bamboo specialists such as the 
critically threatened Purple-winged Dove Claravis godefrida could well 
be present in the area (recently reported not far away from the reserve 
in the Dpto. Canindeyu: Lowen et al. 1995), and would thus also 
benefit from any conservation initiatives that seek to expand the 
reserve’s protected area to the south. 


MARSH SEEDEATER Sporophila palustris (T) 


The species has been erroneously reported at the RNBM (Collar 
et al. 1992, Hayes 1995), the bird in question actually being observed 
at Estancia La Fortuna, Dpto. Canindeyt (P. Scharf in litt. 1994, per 
R. Clay). First records for the RNBM occurred on several dates between 
22 October and 2 November 1994 at Lagunita (24°08’S, 55°25’W), 
where up to three males (part of a mixed Sporophila flock) were 
observed (D. Pullan and R. Denny; also AMN, EZE and R. Clay). 
These dates suggest that this and the following Sporophila seedeaters 
are spring transients (see also Hayes et al. 1994). The first week of 
November also coincides with the arrival on the breeding grounds in 
Corrientes province, Argentina, of this species together with the 
sympatric S. ruficollis, S. hypochroma and S. cinnamomea (Pearman & 
Abadie in press). 


DARK-THROATED SEEDEATER Sporophila ruficollis (NT) 


The species was not previously recorded from Dpto. Canindeyut. 
Records at the RNBM are as follows: two males in cerrado vegetation in 
Aguara Nu at 24°15’S, 55°15’W on 20 October 1994, and one male (part 
of a mixed Sporophila flock; see above) at Lagunita on 29 October 1994 
(AMIN). 


RUFOUS-RUMPED SEEDEATER Sporophila hypochroma (NT) 


The species has not been previously reported for Dpto. Canindeyu. 
Records at the RNBM include at least one male (part of a mixed 
Sporophila flock, see comments under Marsh Seedeater) between 
29 October and 2 November 1994 at Lagunita (AMN, D. Pullan, 
R. Denny, EZE and R. Clay). 


CHESTNUT SEEDEATER Sporophila cinnamomea (NT) 


Records below constitute the first sightings in the RNBM, and the 
second record for the Dpto. Canindeyu: up to three males in a mixed 
Sporophila flock (see comments under Marsh Seedeater) at Lagunita 
between 22 October and 2 November 1994 (D. Pullan, R. Denny, 
AMIN and EZE). 


A. Madroto N. & E. Z. Esquivel ¥/5 Bull. B.O.C. 1997 117(3) 


SOOTY GRASSQUIT Tiaris fuliginosa 

The species was first recorded for Paraguay at the RNBM on 12 
September 1992 (Brooks et al. 1993). Since then, a male was observed 
calling at the same place (24°08'58"S, 55°25’22”W) on 26 September 
1994 (AMN), and another male was seen in November 1994 at 
24°08'03"S, 55°31'44”W (R. Clay). 


Acknowledgements 


We thank R. Clay, F. Hayes, J. Lowen, J. Padwe, M. Pearman and A. Yanosky for 
reading earlier versions of this paper. A final version of this manuscript greatly benefited 
from comments by Nigel Collar. We also want to thank R. Clay, R. Denny, P. Donahue, 
D. Finch, D. Pullan and the “‘Project Yacutinga ’95’’ team members for kindly allowing 
us to cite unpublished observations. The forest rangers of the RNBM have also 
contributed to this paper with their observations. We also want to extend our gratitude to 
The Nature Conservancy (TNC), particularly to Alan Randall for his effort in making 
possible the donation of a GPS receiver to the Fundacion Moisés Bertoni (FMB), and to 
Andrea Cristofani who trained the FMB’s research staff in its proper use. Finally, funds 
to conduct our ongoing research were partly provided by TNC (Adopt an Acre and Parks 
in Peril programmes) and by the FMB’s own resources. Miguel Morales (FMB) deserves 
special credit for providing continuous support to our project. 


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Brooks, T. M., Barnes, R., Bartrina, L., Butchart, S. H. M., Clay, R. P., Esquivel, E. Z., 
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Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
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Hayes, F. E. 1995. Status, distribution and biogeography of the birds of Paraguay. 
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ew York. 


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Hayes, F. E., Scharf, P. A. & Ridgely, R. S. 1994. Austral bird migrants in Paraguay. 
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© British Ornithologists’ Club 1997 


P. Alstrém, U. Olsson & P. Colston 177 Balt B°O.C. 1997 117(3) 


Re-evaluation of the taxonomic status of 
Phylloscopus proregulus kansuensis Meise 


by Per Alstrém, Urban Olsson & Peter R. Colston 
Received 21 August 1996 


Phylloscopus proregulus is generally divided into 3 subspecies: proregulus 
(Pallas) breeding in Siberia, northern Mongolia and northeastern 
China; chloronotus (Gray) in central China and in the Himalayas west to 
central Nepal; and szmlaensis Ticehurst in the westernmost Himalayas 
(e.g. Mayr & Cottrell 1986). Alstr6m & Olsson (1990) argued that 
chloronotus and simlaensis should be treated as specifically different 
from proregulus under the name P. chloronotus, and this has since been 
followed by e.g. Sibley & Monroe (1993) and Beaman (1994). ‘The 
taxon Rkansuensis Meise, described from Lauhukou, northern Gansu 
Province, China (Meise 1933, Stresemann et al. 1937), is either treated 
as a valid subspecies (Ticehurst 1938), a synonym of proregulus (Hartert 
& Steinbacher 1934, Vaurie 1954, Etchécopar & Htie 1983, Meyer de 
Schauensee 1984, Mayr & Cottrell 1986, Williamson 1967) or a 
synonym of chloronotus (Cheng 1987, Alstro6m & Olsson 1990). Based 
on recent field studies of Ransuensis, we propose that it be elevated to 
the rank of species. Throughout this paper, chloronotus refers to the 


subspecies, while P. chloronotus refers to the species (sensu Alstrom & 
Olsson 1990). 


Materials and methods 


On 5 June 1992 Paul Lehman, Francois Vuilleumier and others (Paul 
Lehman in litt.) observed an unidentified Phylloscopus warbler on 
Laoye Shan in the Daban Shan range, Qinghai Province, China 
(36°56'N, 101°40’E; Fig. 1). Tape recordings of the song of this bird 
were sent to P.A. On 31 May-—1 June 1993 P.A., Paul Holt and others 
visited Laoye Shan, where at least 10 singing males of the warbler with 
the unknown song were observed. It was concluded that morphologi- 
cally it appeared to be indistinguishable from P. chloronotus, but both 
song and call were strikingly different from those of chloronotus. 'T'wo of 
these birds were tape recorded (song and calls), and another one was 
caught, measured and photographed, and a blood sample was collected. 
One male was exposed to playback of the songs of proregulus, 
chloronotus and P. sichuanensis (latter described by Alstrém et al. 1992) 
(see Appendix). On 21 June and 4 July U.O. and others found the 
warbler with the unknown song to be common on Huzu Bei Shan in the 
Daban Shan range (c. 37°N, 102°E; Fig. 1). One male was exposed to 
playback of the song of chloronotus (see Appendix). They also observed 
5 males on Laoye Shan on 22 June. One of these was caught, measured 
and photographed, and a blood sample was collected. After consulting 
the literature and specimens (see below), it was concluded that this 
warbler was synonymous with P. proregulus Ransuensis Meise, which 
was collected from much the same area. 


Bull. B.O.C. 1997 117(3) 


178 


P. Alstrém, U. Olsson & P. R. Colston 


P. Alstrém, U. Olsson & P. R. Colston 179 Bull. B.O.C. 1997 117(3) 


On 2-3 June 1994 on Emei Shan, Sichuan Province (29°35'N, 
103°11’E), P.A. exposed 4 territorial, singing males of chloronotus to 
playback of song of Ransuensis (see Appendix). On 7-8 June 1994 P.A. 
found kansuensis to be common (c. 45 individuals) in Xinglong Shan, 
Gansu Province (c. 35°40’N, 103°55’E; Fig. 1). Five of these were 
exposed to playback of the song of proregulus and chloronotus and one to 
P. sichuanensis (see Appendix). On 11-22 June 1994 P.A. surveyed the 
area between Xining, Qinghai Province (36°35’N, 101°55’E; Fig. 1) and 
Jiuzhaigou, Sichuan Province (c. 33°25’N, 104°05’E; Fig. 1), and from 
Jiuzhaigou north to Longxi, Gansu Province (34°59'N, 104°46’'E; 
Fig. 1) in order to try to find out whether or not kansuensis and 
chloronotus were sympatric. There is very little forest in this area 
(except in Jiuzhaigou), and most adequate patches of forest along the 
main road were checked. On 11-14 June 1994 Mengda, Qinghai 
Province (c. 35°45'’N, 102°40’E; Fig. 1) was visited (together with 
Jesper Hornskov), and kansuensis was found to be common (c. 60 
individuals; the commonest bird species). Eight of these were exposed 
to playback of the songs of proregulus and chloronotus (see Appendix), 
and 3 males and 1 female were caught and measured. On 15 June 1994 
4 kansuensis (3 singing males and 1 calling bird, presumably a female) 
were observed in a small patch of forest at Hezuozhen, Gansu Province 
(35°00’, 102°58’E; Fig. 1), and two of the males were exposed to 
playback of proregulus and chloronotus (see Appendix). On 16 June 1994 
chloronotus was found to be fairly common (>13 singing males and 3 
calling birds) in a small forest at Chakou, Gansu Province (c. 34°12'N, 
102°25’E; Fig. 1). Three of these were exposed to playback of 
Ransuensis. No kansuensis were observed at this site. On 17-19 June 
1994 Jiuzhaigou was visited, where several chloronotus but no kansuensis 
were noted. Between Jiuzhaigou and Longxi no suitable forest was 
found. On 22 June 1995 P.A. and P.R.C. visited Laoye Shan, where 
c. 10 Ransuensis were observed. On 23-25 June 1995 P.A. and P.R.C. 
surveyed Huzu Bei Shan, where Rkansuensis was common. On one of 
these a playback test was carried out (see Appendix). 

During the playback experiments a speaker with a 20 m long cable 
was placed in the territory of a singing male. Songs of different taxa 
were played when the bird was considered to be close enough to the 
speaker to hear the song clearly. ‘The term “1st approach’’ is the time 
when the bird exposed to the playback was first seen to move towards 
the speaker. “‘Full response’? means that the bird responded by 
vigorously searching for the source of the sound, while adopting an 
aggressive posture with slightly raised tail and slightly drooped, quickly 
flicking wings; usually silent, but sometimes calling, only rarely singing 


Figure 1. Distribution of chloronotus //// and proregulus (only part of range in Siberia 
shown) \\\\. Detail shows all localities (white figures in black circles) where kansuensis has 
been found: 1, Lauhukou (type locality); 2, Komandse; 3, Hu-dja-dschuang; 4, Laoye 
Shan; 5, Tschau-tou; 6, Huzu Bei Shan; 7, Mengda, 8, Hezuozhen; 9, Xinglong Shan. 
Detail also shows localities (figures in open circles) where chloronotus has been found in 
close proximity to kansuensis: 1, Chakou; 2, Jiuzhaigou. Based on Stresemann et al. (1937) 
and personal observations. 


P. Alstrém, U. Olsson & P. R. Colston 180 Bull. B.O.C. 1997 117(3) 


one or two strophes. The song of P. proregulus was tape recorded by 
P.A. at Changbai Shan, Jilin Province (c 41°30'N, 128°11’E) in June 
1987; the two song types of P. chloronotus were tape recorded by P.A. 
on Emei Shan, Sichuan Province (c. 29°35’N, 103°10’E) in May 1987; 
the song of P. stchuanensis was tape recorded by P.A. in Jiuzhaigou, 
Sichuan Province in June 1989; and the song of Ransuensis was tape 
recorded on Laoye Shan, Qinghai Province in May 1993. 

In the Natural History Museum, Tring, U.K., P.A. and P.R.C. 
examined 1 specimen of kansuensis (collected at " the type locality; 
BMNH 1938.5.16.21) and a further 6 on loan from the Zoologischen 
Museum, Berlin, Germany (collected at or near the type locality; 
including. the holotype), as well as long series of chloronotus and 
proregulus. All of the specimens of Ransuensis and a series of proregulus 
and chloronotus were measured by P.A. Wing length was measured with 
the wing flattened and stretched (maximum chord), and bill length was 
taken to the skull. 


Results 


Vocal differences between kansuensis and proregulus/chloronotus 

The song of kRansuensis begins with a series of faltering, thin, 
high-pitched, slightly harsh tsrip, followed by a row of slightly 
accelerating clear tszp notes (often on two different pitches), and ends in 
a clear c. 1.1-2.2 s long trill (which often changes from high to slightly 
lower pitch) (Fig. 2A). The trill recalls the song of Wood Warbler 
P. sibilatrix and Emei Leaf Warbler P. emeiensis (Alstrom & Olsson 
1995). Sometimes the initial tsvzp notes are omitted, and sometimes the 
trill is not given in every strophe (the song then alternates between ts7ip 
and tsip, the latter often on two pitches, for some time). The song of 
Ransuensis is profoundly different from the varied, somewhat Canary 
Serinus canaria-like song of proregulus (Fig. 3), although the ts7zp notes 
given by kansuensis are somewhat similar to those of proregulus (one 
note marked by an arrow in Fig. 2A and 3, respectively). 

The song of kansuensis is also strikingly different from the two 
different song types of chloronotus (referred to as type A and type B, 
respectively, by Alstr6m & Olsson 1990; Fig. 2B, 2C and 2D). 
However, two of the elements in the repertoire of Ranswensis resemble 
individual elements in chloronotus song. The tsrip notes given by 
Ransuensis are rather close to tsrip notes in chloronotus type B song (one 
marked by an arrow in Fig. 2A, 2B and 2C, respectively), and the tszp 
notes of Ransuensis are rather similar to individual elements in especially 
type B song of chloronotus (one element marked by an * in Fig.2A, 2B 
and 2C, respectively). Accordingly, chloronotus type B song is 


Figure 2. Songs of Ransuensis and chloronotus. Arrows indicate tsrip notes and asterisks 
tsip notes (only one marked in each song). A. Complete song of Ransuensis, Laoye Shan, 
Qinghai, China, June 1993. B. Part of song of chloronotus type B, Emei Shan, Sichuan, 
China, May 1987. C. Part of song of chloronotus, type B (variation), Emei Shan, Sichuan, 
China, June 1994. D. Complete song of chloronotus, type A, Emei Shan, Sichuan, China, 
June 1994. All tape recordings by Per Alstrém. 


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P. Alstrém, U. Olsson & P. R. Colston 183 Bull. B.O.C. 1997 117(3) 


0 1 2 3 Sec 


Figure 4. Calls of Ransuensis, Huzu Bei Shan, June 1995 (A; variation shown); proregulus, 
Huzong, Heilongjiang, China, June 1988 (B); and chloronotus, Emei Shan, Sichuan, 
China, May 1989 (C). Note similarity between calls of proregulus and chloronotus. All tape 
recordings by Per Alstrém. 


somewhat reminiscent of the song of kansuensis when the trills are 
excluded (as is sometimes the case for short periods of time); the most 
striking difference is that the individual tszp elements are double 
(infrequently single, triple or multiple) in chloronotus, while they are 
single in Ransuensis. 

The call of Ransuensis is a thin tsi-di or tst-di-di (Fig. 4A); sometimes 
it consists of four or five syllables, tst-di-di-di or tst-di-di-di-di, and 
rarely it is a monosyllabic tszt or, differently transcribed, tsiit (Fig. 4A). 
It is significantly different from the soft, subdued dju-ee or duee 
of proregulus (Fig. 4B) and the monosyllabic tswist or, differently 
transcribed, uzst of chloronotus (Fig. 4C). 


Playback tests 

Eleven of the 17 (65%) Ransuensis exposed to playback of proregulus 
showed no interest whatsoever in the song of proregulus, while 6 
individuals (No.5, 7, 8, 11, 12 and 13 in Appendix) reacted to the song 
of proregulus. However, in three of the individuals which did react to 
the song of proregulus (No. 7, 12 and 13) there was no aggression at all, 
the birds only showed a very temporary interest, which was interpreted 
as merely curiosity (see Appendix). In two others (No. 8 and 11) the 
aggression towards the song of proregulus was not nearly so strong as to 
the song of Ransuensis, and it ceased after a while (see Appendix). Only 
one kansuensis (No. 5) responded with strong aggression towards the 
song of proregulus, though it did diminish after some time (see 
Appendix). It should be noted that four (No. 5, 7, 8 and 11) of the 
Ransuensis which reacted towards the song of proregulus also reacted 
towards the song of chloronotus (see below). 

Thirteen out of the 18 (72%) Ransuensis exposed to playback of the 
song (both types) of chloronotus did not respond at all to chloronotus 
song. Five (No. 5, 7, 8, 11 and 12) individuals responded to chloronotus 
type B song. However, in two of these (No. 5 and 12) there was no 


P. Alstrém, U. Olsson & P. R. Colston 184 Bull. B.O.C. 1997 117(3) 


TABLE 1 
Measurements of chloronotus (from China and NE India), Ransuensis and proregulus (from 
Siberia and S China) based on personal measurements of specimens in The Natural 
History Museum, Tring, U.K., specimens on loan from the Zoologischen Museum, 
Berlin, Germany, and live birds. Numbers in brackets refer to means and standard 
deviations. All measurements in mm 


male female 
chloronotus wing 50.5-57.0 (54.1; 1.85) 48.0-51.5 (49.7; 1.06) 
(n=14 males, 10 females) tail 36.0-44.5 (39.9; 2.13) 33.5-38.5 (36.5; 1.73) 
bill 9.8-11.4 (10.6; 0.48) 10.1-10.9 (10.4; 0.27) 
Ransuensis wing 54.0-57.5 (55.6; 1.11) 51.0—52.0 (51.5; 0.35) 
(n=9 males, 5 females) tail 40.0-44.0 (42.4; 1.45) 37.0—40.0 (38.4; 1.29) 
bill 10.5-11.3 (10.8; 0.24) 10.0-11.1 (10.5; 0.41) 
proregulus wing 49.0—-54.5 (51.3; 1.91) 48.0—52.0 (49.8; 1.37) 
(n=12 males, 14 females) tail 34.5-39.5 (36.7; 1.70) 33.0—40.0 (36.2; 1.95) 
bill 9.7-10.7 (10.3; 0.34) 9.7-10.8 (10.3; 0.33) 


aggression at all involved, and the reaction was interpreted as merely 
curiosity. In none of the others was the reaction to the song of 
chloronotus nearly so strong as to the song of Ransuensis, and the interest 
in the song of chloronotus invariably ceased after some time (see 
Appendix). Only one kansuensis (No. 7) reacted to chloronotus type A 
song, though there was no apparent aggression involved. 

None of the 7 chloronotus tested with the song of Ransuensis showed 
any aggression towards this song, though individual number 3 showed 
temporary interest the third time it was exposed to Ransuensis song (see 
Appendix). 


Morphological differences between kansuensis and proregulus/ 
chloronotus 

Kansuensis differs from proregulus mainly in being clearly paler 
yellow on the supercilium (unless very worn, proregulus is bright yellow 
on especially the anterior part of the supercilium, while Ransuensis 
shows only a very faint yellowish tinge to the supercilium in front 
of/above the eye). At least in spring and summer the lower mandible is 
generally paler in Ransuensis than in proregulus: it is either entirely pale 
orange or pale orange with a very small dark tip in Ransuensis, while it 
has a much more extensive dark tip in proregulus (lower mandible 
frequently appears nearly all dark, although it is sometimes extensively 
pale orange or even practically all pale orange). Also the legs generally 
appear paler in Ransuensis than in proregulus, although there is overlap. 
Furthermore, kansuensis has significantly longer wings (Table 1; 
Mann-Whitney U test, P..43-<=0-0002, Promales= 0-01) and tail (Table 1; 
Mann-Whitney U test, P..4;.,=0-0001, Pronales= 0-04) and a different 
wing formula (Table 2). . 

Compared to chloronotus, there appears to be a tendency for the 
supercilium to be marginally more yellowish-tinged, the lateral 


ales 


P. Alstrém, U. Olsson & P. R. Colston 185 Bull. B.O.C. 1997 117(3) 


TABLE 2 
Wing formulae of chloronotus, Ransuensis and proregulus. Based on same specimens as in 
Table 1 (both sexes combined). Wp means wing-point and P means primary. P10 is 
compared to tips of primary coverts, other primaries to wing-point. Figures given are 
mean, range and standard deviation 


chloronotus Ransuensis proregulus 

wp: P7 13.3% 15.4% 87.5% 

wp: P6 6.7% 46.1% 0% 

wp: P7=6 80.0% 38.5% 12.5% 

P10 + $.0.(5.5—9.5- 1.19) + 7.6 (5.5—9.0; 0.97) + 6.9 (4.5—9.0; 1.39) 
P9 — 8.6 (6.5-11.0; 1.53) — 8.1 (6.5-10. 0: 1.10) — 6.7 (5.0—8.0; 0.83) 
P8 — 1.4 (0.5-2.0; 0.48) — 1.4 (1.0-2.0; 0. 34) ='0F-(0.5+1-5;°0.32) 
P5 — 1.3 (0.5-3.0; 0:72) — 1.3 (0.5-2.0; 0.48) — 1.9 (1.0-3.5; 0.64) 


crown-stripes marginally paler and greener, and the underside whiter 
in Ransuensis, but these differences are so subtle that Ransuensis and 
chloronotus are essentially identical on plumage. However, the lower 
mandible is generally clearly paler in Ransuensis than in chloronotus (in 
the latter it frequently appears nearly all dark, although sometimes pale 
orange with a very small dark tip). Also the legs generally appear paler 
in Ransuensis than in chloronotus, although there is overlap. Although 
Ransuensis and chloronotus are basically very similar on measurements 


and wing formulae, Ransuensis has marginally longer wings (Table 1; 
Mann-Whitney U test, P =0.045, Pronales 0-006) and a greater 


males 


tendency for the 6th primary to be equal to the 7th (‘Table 2). 


Breeding habitat of kansuensis, proregulus and chloronotus 

At Laoye Shan (altitude c. 2500-2900m) and Mengda (ce. 
2200-2500 m) kansuensis occurs in predominantly deciduous forest 
(including e.g. birch Betula and aspen Populus) with some spruce Picea 
mixed in (overall <10-c. 20%). On Huzu Bei Shan (c. 2700-2900 m) it 
occurs mainly in deciduous forest (predominantly birch) with some 
spruce and tall junipers Juniperus mixed in, much less commonly in 
predominantly coniferous forest. At Xinglong Shan kansuensis occurs 
mainly in mixed deciduous and spruce forest (the predominant forest 
type), but also in mainly deciduous as well as mainly spruce forest, at 
an altitude of c. 1700-1800 m. At Hezuozhen kansuensis was found in 
“semi-old’’ secondary spruce forest with much undergrowth of 
deciduous bushes at an altitude of c. 3200m (altitude according to 
locals). 

The breeding habitats of proregulus and chloronotus differ signifi- 
cantly from that favoured by kansuensis. Proregulus breeds in the taiga, 
in coniferous forest or mixed forest with a high percentage of conifers 
(Dement’ev & Gladkov 1954, Flint et al. 1984, Rogacheva 1992, 
pers. obs.). Chloronotus breeds chiefly in spruce/fir Abies forest or 
predominantly spruce/fir forest, and only very sparsely in mainly 
deciduous forest (on mountains, just below the spruce forest belt). In 


P. Alstrém, U. Olsson & P. R. Colston 186 Bull. B.O.C. 1997 117(3) 


China chloronotus breeds between c. 2000 and c. 4000 m, usually at c. 
2600—c. 3100 m (Etchécopar & Htie 1983, Meyer de Schauensee 1984, 
Alstrom et al. 1992, pers. obs.). 


Breeding ranges of kansuensis, proregulus and chloronotus 

Kansuensis has been observed in the breeding season at 9 localities, 
from the eastern Lenglong Ling, north Gansu Province (c. 37°30'N, 
102°30’E) in the north to Hezuozhen, south Gansu in the south (Fig. 1). 
It seems likely that its range extends at least slightly further northwest, 
as the mountain range continues in that direction. It is not known 
where Rkansuensis winters, but due to the severe winter climate in its 
breeding range, it ought to be considerably further south. In 1993 
Ransuensis apparently left Laoye Shan in mid to late October Jesper 
Hornskov 2n Iitt.). 

The breeding range of proregulus appears to be disjunct from that of 
Ransuensis by at least 1000 km (Fig. 1). Mayr & Cottrell (1986) and 
Cheng (1987) state that proregulus and chloronotus intergrade in eastern 
Qinghai. This surely refers to Ransuensis. We have found no evidence of 
proregulus breeding in Qinghai. 

Chloronotus (including simlaensis) breeds from the western Himalayas 
through central China north to at least Chakou (Fig. 1), at the most 
100 km south of Hezuozhen, where kansuensis was found. It seems 
likely that the breeding ranges of Ransuensis and chloronotus actually 
overlap marginally, although this has not yet been proven. 


Discussion 


Since Ransuensis is morphologically more similar to chloronotus than to 
proregulus, it may seem surprising that most previous authors (Hartert 
& Steinbacher 1934, Vaurie 1954, Etchécopar & Hiie 1983, Meyer de 
Schauensee 1984, Mayr & Cottrell 1986, Williamson 1967) have 
lumped kansuensis with proregulus rather than with chloronotus. 
However, Hartert & Steinbacher (op. cit.) do not state how many 
individuals they studied, Vaurie (op. cit.) only examined one, and we 
doubt that any of the others actually examined specimens of kansuensis. 

The morphological differences between Ransuensis and chloronotus are 
so slight that, based on these alone, kansuensis would be _ best 
synonymized with chloronotus or considered a very poorly differentiated 
subspecies of P. chloronotus. In contrast, the vocalizations of Ransuensis 
are very different from those of chloronotus. In fact, the differences in 
song between kansuensis and chloronotus are much more pronounced 
than between different species in some other presumably monophyletic 
groups of Phylloscopus warblers, e.g. P. occipitalis-P. reguloides- 
P. davisoni (Martens 1980, Alstro6m & Olsson 1993), P. schwarzi-P. 
armandiu (Martens 1980, Alstrom & Olsson 1994), and P. griseolus- 
P. affinis-P. subaffinis (Martens 1980, Alstr6m & Olsson 1992, 1994), 
and at least as pronounced as between other species of Phylloscopus. 
This alone suggests that the rank of species would be appropriate for 
Ransuensis. However, since chloronotus has two song types which are 
nearly as different from each other as from the song of Rkansuensis, the 


P. Alstrém, U. Olsson & P.R. Colston 187 Bull. B.O.C. 1997 117(3) 


distinctive song of kansuensis might be considered to be just a third, 
geographically localized, variant of P. chloronotus song. ‘This is 
contradicted by the playback tests which have been carried out, which 
instead indicate that the songs of kansuensis and chloronotus would act as 
prezygotic reproductive isolating mechanisms if there were any 
sympatry. Especially the playback tests on those Ransuensis (No. 16 and 
17) and chloronotus (No. 5, 6 and 7) which were found in close 
proximity to each other (separated by at the most 100 km), combined 
with the apparent lack of intergradation between these two taxa (as 
indicated by the lack of individuals with intermediate vocalizations ), 
strongly suggest that Ransuensis and chloronotus should be considered 
separate species. It is curious that 5 of the Rkansuensis tested showed 
some interest (though there was no or relatively little aggression 
involved) in the type B song of chloronotus, while only one individual 
reacted with curiosity to chloronotus type A song. Since chloronotus 
reacts equally strongly to both of its two song types (Alstroém & Olsson 
1990 and Appendix), the reason why Ransuensis showed more interest in 
the type B song than in the type A song does not seem to be a case of 
the former song type being more important in territory defence than 
the latter. It seems possible that Ranswensis considers the type B song to 
be more reminiscent of its own song than the type A song. In general, 
response from one taxon to playback of song of another taxon is of little 
taxonomic relevance. Response to playback of heterospecific closely 
related sympatric taxa has been noted in several cases, presumably 
because of interspecific territorialism (e.g. Emlen et al. 1975, Catchpole 
1978, Catchpole & Leisler 1986, Prescott 1987, Elfstro6m 1990, Baker 
1991). Response to playback of allopatric taxa is equally uninformative 
in this context, and may simply be a result of similarities between the 
songs of the taxa involved (cf. Ratcliffe & Grant 1985); the song’s 
function as a reproductive isolating barrier is unlikely to be fully 
developed if the taxa are geographically separated. The fact that 
kansuensis and chloronotus exist so close to each other without any signs 
of intergradation indicates that they have evolved independently of 
each other for a substantial period of time. Significant interbreeding 
would presumably have merged the two forms. The differences in 
breeding habitat are further evidence of speciation (Richman & Price 
1992). 

The overall similarity between kansuensis, P. chloronotus and P. 
proregulus suggests that they share a common ancestor and thus form 
a monophyletic group. On plumage, wing-formula, size and song 
kansuensis shows a greater similarity to chloronotus than to proregulus. 


"We assume that the offspring from any mixed pairs of kansuensis and chloronotus 
would have aberrant songs compared to their parent taxa. This assumption is supported 
by reports of aberrant songs in suspected hybrids between Phylloscopus bonelli x P. 
sibilatrix (Bremond 1972, Fouarge 1972) and P. trochilus x P. collybita (Da Prato & Da 
Prato 1986). However, since song appears to be to a great extent learned in “‘song-birds”’ 
in general (see review in Catchpole & Slater 1995), it is possible that the song of hybrids 
would be very similar to or indistinguishable from the species which is more numerous in 
the area where it was born. 


P. Alstrém, U. Olsson & P. R. Colston 188 Bull. B.O.C. 1997 117(3) 


This, together with the distributional pattern, suggests that Ransuensis 
and chloronotus diverged more recently, and thus are more closely 
related to each other than to proregulus. In analogy with the proposed 
treatment of kansuensis and chloronotus as separate species, Ransuensis 
and proregulus must also be treated as specifically different. The 
playback tests support this treatment. However, three Ransuensis (No. 
5, 8 and 11) reacted with some aggression toward the song of proregulus, 
and in one of these (No. 5) the response was almost as strong as to the 
song of kansuensis. It should be noted that these three birds also 
responded to chloronotus type B song. As discussed above, it is 
important to keep in mind that only absence of response to playback 
may have some taxonomic relevance. The differences in breeding 
habitat between Ransuensis and proregulus further support the view that 
they are better treated as separate species. 

It is clear that Ransuensis is not conspecific with P. sichuanensis. 
These two were found in sympatry at Laoye Shan, Xinglong Shan, 
Mengda, Hezuozhen and Chakou, and morphologically and vocally 
they are significantly different (Alstr6m et al. 1992). Also, the two 
kansuensis (No. 1 and 3) which were exposed to playback of the song of 
P. sichuanensis did not respond at all to it. Moreover, where both taxa 
occurred together, there was a difference in average habitat preference, 
sichuanensis favouring less-tall secondary growth at lower altitude than 
Ransuensis. 

P. proregulus (sensu lato) has been variously named Pallas’s Warbler, 
Pallas’s Leaf Warbler, Pallas’s Willow Warbler, Lemon-rumped 
Warbler and Pale-rumped Warbler. We support Beaman (1994) in 
using the name Pallas’s Leaf Warbler for P. proregulus (sensu stricto), 
Lemon-rumped Warbler for P. chloronotus (sensu Alstro6m & Olsson 
1990), and suggest the name Gansu Leaf Warbler for P. Ransuensis. 
There are two reasons why we prefer the name Gansu Leaf Warbler 
rather than “‘Qinghai Leaf Warbler’ (which might be thought a more 
suitable name, since nearly all of the records of kansuensis are from 
Qinghai Province and only a few from Gansu Province): firstly, the 
name Gansu Leaf Warbler is a translation of the scientific name 
(Gansu is the modern spelling of Kansu), and, secondly, the name 
Qinghai would surely be mis-pronounced by most people (correct 
pronunciation “‘Chinghigh’’). 


Summary 


Phylloscopus proregulus kansuensis Meise has variously been treated as a distinct 
subspecies, a synonym of P. chloronotus (proregulus) chloronotus or a synonym of P. (p.) 
proregulus (most authors). It is morphologically only very slightly different from 
chloronotus, though more clearly separable from proregulus (especially by its much paler 
yellow supercilium). Both song and calls are strikingly different from those of both 
chloronotus and proregulus (most different from latter). Unlike chloronotus and proregulus it 
breeds mainly in deciduous or mixed forest. In the breeding season it is parapatric with 
chloronotus (without any known geographical overlap), while it appears to be widely 
allopatric with proregulus. Playback tests indicate that the songs would act as prezygotic 
reproductive isolation mechanisms if there were any sympatry. We suggest that 
kansuensis be treated as a distinct species and that the English name be Gansu Leaf 
Warbler. 


P. Alstrém, U. Olsson & P. R. Colston 189 Bull. B.O.C. 1997 117(3) 


Acknowledgements 


Thanks to Paul Lehman for sending a tape to P.A., thereby drawing his attention to the 
existence of Ransuensis. We are grateful to Dr W. Meise, Dr Jochen Martens and Dr 
Robert Prys-Jones for their valuable comments on a draft of the manuscript. P.A. is 
grateful to Paul Holt, Jesper Hornskov, Delores Jensen, Richard Newton, Bob Sharland, 
Dr Bob Wilson, Dr John K. Wilson and Dr Joseph Wilson for assistance in the field. 
Thanks to Mark Beaman for comments on the English name of kansuensis, and to Matti 
Ahlund for help with statistics. We are also grateful to the Zoologischen Museum, Berlin, 
Germany for lending us specimens of Ransuensis and to The Natural History Museum, 
Tring, U.K. for granting us access to its collection. 


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Addresses: Per Alstr6m and Urban Olsson, University of Géteborg, Dept. of Zool., Sect. 
of Morphology and Systematics, Medicinaregatan 18, 413 90 Géteborg, Sweden. 
Peter R. Colston, The Bird Group, The Natural History Museum, Akeman Street, 
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© British Ornithologists’ Club 1997 


Appendix 
Playback experiment data 
Ransuensis 


Individual No. 1, Laoye Shan 31 May 1993 

e proregulus (2 min). No response. e sichuanensis (2 min). No response. e chloronotus type 
A (2 min). No response. e chloronotus type B (2 min). No response. e Ransuensis (2 min). 
Full response. e proregulus (2 min). No response. e sichuanensis (2 min). No response. 
e chloronotus type A (2 min). No response. e chloronotus type B (2 min). No response. 
e Ransuensis (2 min). Full response. 


Individual No. 2, Huzu Bei Shan 21 June 1993 
e chloronotus type A (2 min). No response. e kansuensis (2 min). Full response. 


Individual No. 3, Xinglong Shan 8 June 1994 

e proregulus (2 min). No response. e sichuanensis (2 min). No response. e chloronotus type 
A (2 min). No response. e chloronotus type B (2 min). No response. e kansuensis (2 min). 
1st approach at 5 s followed by full response rest of time. e chloronotus type B (4 min). No 
response. e chloronotus type A (4 min). No response. e proregulus (4 min). No response. 
e kansuensis (2 min). 1st approach at 8s followed by full response rest of time. 
e sichuanensis (2 min). No response. e chloronotus type B (2 min). No response. 
e chloronotus type A (2 min). No response. e kansuensis (2 min). 1st approach at 9s 
followed by full response rest of time. e proregulus (2 min). No response. e kansuensis 
(2 min). 1st approach at 4s followed by full response rest of time. 


Individual No. 4, Xinglong Shan 8 June 1994 

e proregulus (2 min). No response. e kansuensis (2 min). 1st approach at 8 s followed by 
full response rest of time. e chloronotus type A (2 min). No response. e chloronotus type B 
(2 min). No response. e kansuensis (2 min). 1st approach at 5 s followed by full response 
rest of time. e chloronotus type B (2 min). No response. e kansuensis (2 min). 1st approach 
18s followed by full response rest of time. e chloronotus type A (2 min). No response. 
e Ransuensis (2 min). 1st approach at 110s followed by full response rest of time. 


P. Alstrém, U. Olsson & P. R. Colston 191 Bull. B.O.C. 1997 117(3) 


Individual No. 5, Xinglong Shan 8 June 1994 

e chloronotus type B (2 min). No response. e kansuensis (2 min). 1st approach at 7s 
followed by full response rest of time. e chloronotus type B (2 min). No response. 
e kansuensis (2 min). 1st approach at 3s followed by full response rest of time. 
e chloronotus type A (2 min). Came close to speaker twice (at 18 s and 42 s), but showed 
no aggression. e kansuensis (2 min). 1st approach at 4s followed by full response rest 
of time. e proregulus (2 min). 1st approach at 4s followed by full response rest of time. 
e proregulus (after the speaker had been moved c. 20 m; 4 min). 1st approach 3s. Less 
strong response than before, on and off during rest of time. e chloronotus type A (2 min). 
No response. 


Individual No. 6, Xinglong Shan 8 June 1994 

e chloronotus type B (2 min). No response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e kansuensis (2 min). 1st approach at 6s 
followed by full response rest of time. e proregulus (2 min). No response. e chloronotus 
type A (2 min). No response. e kansuensis (2 min). 1st approach at 5s followed by full 
response rest of time. e chloronotus type B (2 min). No response. 


Individual No. 7, Xinglong Shan 8 June 1995 

e proregulus (4 min). Came to c. 3m from speaker at 25s, but moved away at c. 35s. 
Showed no aggression. No further response. e chloronotus type A (4 min). 1st approach at 
6s. Appeared curious, not aggressive. At 36s c. 3m from speaker. Moved away after 
that. e chloronotus type B (4 min). Ist approach at 18s. At 50s c. 3m from speaker. 
Remained close to speaker rest of time; appeared slightly annoyed. e Ransuensis (2 min). 
ist approach at 8 s followed by full response rest of time. Much more agitated than when 
proregulus and the two types of chloronotus were played. e proregulus (2 min). No 
response. e chloronotus type B (2 min). 1st approach at 9s, but no further response. 
e kansuensis (2 min). 1st approach at 7 s followed by full response rest of time. 


Individual No. 8, Mengda 12 June 1994 

e proregulus (2 min). No response. e chloronotus type A (2 min). No response. 
e kansuensis (2 min). 1st approach at 10s followed by full response rest of time. 
e chloronotus type B (4 min). 1st approach at 5s followed by full response until c. 25s, 
thereafter gradually turning uninterested, and after c. 1 min no response at all. 
e proregulus (4 min). 1st approach at 28s followed by full response for c. 1 min, then 
gradually less interested. e chloronotus type A (4 min). No response. e chloronotus type B 
(4 min). No response. e kansuensis (2 min). 1st approach at 7 s followed by full response 
rest of time. 


Individual No. 9, Mengda 12 June 1994 

Ransuensis ( a few s). Immediately full response. e proregulus (2 min). No response. 
chloronotus type A (2 min). No response. e chloronotus type B (2 min). No response. 
Ransuensis (2 min). 1st approach at 18s followed by full response rest of time. 
proregulus (2 min). No response. e chloronotus type A (2 min). No response. 
kansuensis (2 min). 1st approach at 4s followed by full response rest of time. 
chloronotus type B (2 min). No response. e kansuensis (2 min). 1st approach at 27s 
followed by relatively weak response rest of time. 


Individual No. 10, Mengda 12 June 1994 

e Rkansuensis (a few s). Immediately full response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e chloronotus type B (2 min). No response. 
e kansuensis (2 min). 1st approach at 14s followed by full response rest of time. 
e@ proregulus (2 min). No response. e chloronotus type A (2 min). No response. 
e chloronotus type B (2 min). No response. e kansuensis (2 min). Relatively weak response. 


Individual No. 11, Mengda 12 June 1994 

e kansuensis (c. 30s). Immediately full response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e chloronotus type B (4 min). 1st approach at 
24 s. Some response; approached speaker, flicked wings now and then. At c. 2 min 10s it 
moved away. Response interpreted as mainly curiosity. e Ransuensis (2 min). 1st approach 
at 6 s followed by full response rest of time. Much stronger response than to chloronotus. 
e proregulus (4 min). 1st approach at c. 20s. Some response; approached speaker, flicked 
wings now and then until c. 2 min, when it moved away. Response interpreted as mainly 
curiosity. e chloronotus type A (4 min). No response. e chloronotus type B (4 min). 1st 


P. Alstrém, U. Olsson & P. R. Colston 192 Bull. B.O.C. 1997 117(3) 


approach at 18s. Obviously interested in the song; searched for the source of the sound 
and flicked its wings now and then. At c. 1 min 50s it moved away from speaker. 
e Rkansuensis (2 min). 1st approach at 3s followed by full response rest of time. Much 
stronger response than to proregulus and chloronotus. 


Individual No. 12, Mengda 12 June 1994 

e kansuensis (a few s). Immediately full response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e chloronotus type B (4 min). 1st approach at 
c. 15s, but no further response. e Rkansuensis (2 min). 1st approach at 45 s (bird was not 
seen when playback started, so perhaps had moved out of hearing range) followed by full 
response rest of time. e proregulus (4 min). Moved somewhat closer to speaker, but was at 
the most curious, definitely not aggressive. e chloronotus type A (4 min). No response. 
e chloronotus type B (4 min). No response. e kansuensis (2 min). 1st approach at 4s 
followed by full response rest of time. 


Individual No. 13, Mengda 12 June 1994 

e@ kansuensis (a few s). Immediately full response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e chloronotus type B (2 min). No response. 
e kansuensis (2 min). 1st approach at 3s followed by full response rest of time. 
e proregulus (2 min). 1st approach at 6s, but no further response. e chloronotus type A 
(2 min). No response. e chloronotus type B (2 min). No response. e Ransuensis (2 min). 1st 
approach at 3s followed by full response rest of time. 


Individual No. 14, Mengda 13 June 1994 

e proregulus (2 min). No response. e chloronotus type A (2 min). No response. 
e chloronotus type B (2 min). No response. e Ransuensis (2 min). 1st approach at 12s 
followed by full response rest of time. e proregulus (2 min). No response. e chloronotus 
type A (2 min). No response. e chloronotus type B (2 min). No response. e kansuensis 
(2 min). 1st approach at 8s followed by full response rest of time. 


Individual No. 15, Mengda 14 June 1994 

e kansuensis (a few s). Immediately full response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e kansuensis (c. 30s). Immediately full 
response. e chloronotus type B (2 min). No response. e Ransuensis (1 min). 1st approach at 
8 s followed by full response rest of time. e proregulus (2 min). No response. @ Ransuensis 
(c. 30s). Immediately full response. e chloronotus type A (2 min). No response. 
e chloronotus type B (2 min). No response. e kansuensis (2 min). 1st approach at 4s 
followed by full response rest of time. e chloronotus type B (2 min). No response. 


Individual No. 16, Hezuozhen 15 June 1994 

e kansuensis (a few s). Immediately full response. e proregulus (2 min). No response. 
e chloronotus type A (2 min). No response. e chloronotus type B (2 min). No response. 
e kansuensis (2 min). 1st approach at 18s followed by full response rest of time, but not 
quite so aggressive as usual. e proregulus (2 min). No response. e chloronotus type A 
(2 min). No response. e chloronotus type B (2 min). No response. e kansuensis (2 min). 1st 
approach at 14s followed by full response rest of time, but not quite so aggressive as 
usual. 


Individual No. 17, Hezuozhen 15 June 1994 

e kansuensis (a few s). Immediately full response. e chloronotus type B (2 min). No 
response. e kansuensis (a few s). Immediately full response. e chloronotus type A (2 min). 
No response. e kansuensis (a few s). Immediately full response. e proregulus (2 min). No 
response. e kansuensis (a few s). Immediately full response. e chloronotus type B (2 min). 
No response. @ chloronotus type A (2 min). No response. e kansuensis (2 min). 1st 
approach at 8 s followed by full response rest of time. e proregulus (2 min). No response. 
e kansuensis (a few s). Immediately full response. e chloronotus type A (2 min). No 
response. @ kansuensis (a few s). Immediately full response. e chloronotus type B (2 min). 
No response. e kansuensis (1 min). 1st approach at 6s followed by full response rest of 
time. @ chloronotus type B (2 min). No response. 


Individual No. 18, Huzu Bei Shan 24 June 1995 

e chloronotus type A (4 min). No response. e kansuensis (1 min). 1st approach at 35s 
followed by full response rest of time. e chloronotus type A (4 min). No response. 
e kansuensis (1 min). 1st approach at 12s followed by full response rest of time. 


P. Alstrém, U. Olsson & P. R. Colston 193 Bull. B.O.C. 1997 117(3) 


e chloronotus type B (2 min). No response. e Ransuensis (1 min). 1st approach at 6s 
followed by full response rest of time. e chloronotus type B (2 min). No response. 


chloronotus 


Individual No. 1, Emei Shan 2 June 1994 

e kansuensis (2 min). No response. e chloronotus (2 min). 1st approach at 16s followed by 
full response rest of time. e kansuensis (2 min). No response. e chloronotus (2 min). 1st 
approach at 6s followed by full response rest of time. e kansuensis (2 min). No response. 


Individual No. 2, Emei Shan 3 June 1994 

e Ransuensis (2 min). No response. e chloronotus type A (2 min). 1st approach at 4s 
followed by full response rest of time. e kansuensis (2 min). No response. e chloronotus 
type A (2 min). Ist approach at 5s followed by full response rest of time. e Ransuensis 
(2 min). No response. e chloronotus type B (2 min). 1st approach at 6s followed by full 
response rest of time. 


Individual No. 3, Emei Shan 3 June 1994 

e proregulus (2 min). No response. e sichuanensis (2 min). No response. e Rkansuensis 
(4 min). No response. e chloronotus type B (2 min). 1st approach at 7s followed by full 
response rest of time. e proregulus (2 min). No response. e sichuanensis (2 min). No 
response. e kansuensis (4 min). No response. e chloronotus type B (2 min). 1st approach at 
4s followed by full response rest of time. e Ransuensis (4 min). No response, though at 
35s and 3 min 40s came close to speaker, but showed no aggression, and moved off 
almost immediately. e chloronotus type B (2 min). Ist approach at 7s followed by full 
response rest of time. 


Individual No. 4, Emei Shan 3 June 1994 

e Ransuensis (2 min). No response. e chloronotus (2 min). 1st approach at 8 s followed by 
full response rest of time. e Ransuensis (2 min). No response. @ chloronotus type A (2 min). 
1st approach at 4s followed by full response rest of time. 


Individual No. 5, Chakou 16 June 1994 

e chloronotus (a few s). Immediately full response. e Ransuensis (2 min). No response. 
e chloronotus type B (1 min). 1st approach at 12 s followed by full response rest of time. 
e kansuensis (2 min). No response. e chloronotus type A (2 min). 1st approach at 9s 
followed by full response rest of time. e Ransuensis (2 min). No response. e chloronotus 
type B (a few s). Immediately full response. 


Individual No. 6, Chakou 16 June 1994 

e chloronotus (a few s). Immediately full response. e kansuensis (2 min). No response. 
e chloronotus type B (2 min). 1st approach at 8s followed by full response rest of time. 
e Rkansuensis (2 min). No response. e chloronotus type A (2 min). 1st approach at 4s 
followed by full response rest of time. 


Individual No. 7, Chakou 16 June 1994 

e chloronotus (a few s). Immediately full response. e kansuensis (2 min). No response. 
e chloronotus type-A (2 min). 1st approach at 14s followed by full response rest of time. 
e kansuensis (2 min). No response. e chloronotus type A (2 min). 1st approach at 9s 
followed by full response rest of time. e Ransuensis (2 min). No response. e chloronotus 
type A (2 min). Ist approach at 4s followed by full response rest of time. e Ransuensis 
(2 min). No response. e chloronotus type B (2 min). 1st approach at 4s followed by full 
response rest of time. 


G. F. Barrowclough et al. 194 Bull. B.O.C. 1997 117(3) 


New records of birds from Auyan-tepui, 
Estado Bolivar, Venezuela 


by George F. Barrowclough, Miguel Lentino R. & 
Paul R. Sweet 


Received 6 Fuly 1996 


Auyan-tepui, located in Estado Bolivar, southern Venezuela, is one of 
the largest of the isolated sandstone mesas or tepuis of the highland 
region of south Venezuela that has come to be known as the Pantepui 
(Mayr & Phelps 1967). Auyan-tepui was first explored ornithologically 
during the Phelps Venezuela expedition of 1937-1938 (Chapman 1939, 
Gilliard 1941). That expedition spent several months on the southern 
talus slopes of the mountain, but due to arduous conditions and logistic 
difficulties, was only able to explore a small area on the southern, drier 
end of the 700 square kilometre summit plateau. Despite this, 32 
species were recorded on the plateau including several new subspecies. 
Mayr & Phelps (1967) considered this an adequate sample for 
comparative studies of the avifauna. Subsequent to the 1937 
expedition, the Universidad Central de Venezuela organised a brief 
expedition in 1956 that spent approximately one month on the extreme 
south end of the plateau and obtained a few specimens of birds as part 
of a general collection that emphasised plants (Steyermark 1967); the 
ornithological results of the 1956 expedition have not been reported in 
the literature. The geology, geomorphology and botany of the tepui 
have been reviewed by Tate (1938) and Berry et al. (1995). 

As part of the recent Robert G. Goelet American Museum—Terramar 
Expedition to Auyan-tepui, we collected birds on the summit from 
1 February to 1 March 1994. Five helicopter-supported camps were 
established in widely dispersed areas of the tepui summit, enabling us 
to sample the avifauna from a variety of habitats and elevations. Based 
on our Global Positioning System estimates (Appendix 1) of latitude 
and longitude, modern side-scanning radar images of Auyan-tepui, and 
the published map from the 1937/38 expedition (Tate 1938), it appears 
that our camps I and V were very close to the Phelps Expedition’s 
1850 m and 2200 m camps. Our other three camps sampled areas not 
previously visited by ornithologists, including a very wet area just 
above Angel Falls. Specimens were obtained using mistnets and 
shotguns and are deposited in the collections of the American Museum 
of Natural History (AMNH), New York, the Estacion Biologica de 
Rancho Grande (EBRG), Maracay, Venezuela, and the Coleccion: 
Ornitologica Phelps (COP), Caracas. 

In this paper, we report 21 new species records for the summit 
plateau of Auyan-tepui; this includes one first record collected by the 
1956 expedition from the Universidad Central de Venezuela. Several of 
our new records can be attributed to our use of mistnets, a sampling 
technique not used on early expeditions to localities in the Pantepui. 
The use of mistnets explains our collection of the Aegolius owl, the 


G. F. Barrowclough et al. 195 Bull. B.O.C. 1997 117(3) 


oilbird, and several new hummingbirds. In one calendar month (four 
man months), we added 21 new species to a summit avifauna that had 
numbered 32. We continue to believe (e.g. Barrowclough et al. 1995) 
that the avifaunas of most elements of the Venezuelan Pantepui are 
inadequately known due to restricted sampling duration, seasons, and 
techniques. A complete list of birds known to date from the summit 
plateau of Auyan-tepui is included in Appendix 2. 


New records for Auyan-tepui 
TEPUI TINAMOU Crypturellus ptaritepui 


This tinamou, described in 1945, was previously known from only 
six specimens collected on Ptari-tepui and adjacent Sororopan-tepui 
(Phelps & Phelps 1958); those tepuis are some 50 km distant from 
Auyan-tepui and separated from it by the elevated (500-1000 m) 
grassland known as La Gran Sabana. The tinamou had been recorded 
as occurring in cloud forest in the subtropical zone between 1350 and 
1800 m (Meyer de Schauensee & Phelps 1978). On Auyan-tepui four 
specimens were taken at three localities, from approximately 1500 to 
1700 m, in the more humid northern areas of the tepui. The species was 
encountered frequently both in forested and more open scrubby areas 
but was retiring in its habits and usually was seen running into cover. 
One of two males collected had moderately enlarged testes and one of 
two females had an enlarged ovary; none were in moult. Vocalisations 
of an unknown tinamou at high elevations on Chimanta-tepui (c. 50 km 
south of Auyan-tepui) were tentatively assigned to this species by 
Medina Cuervo (1992). The failure of the 1937/38 Phelps expedition to 
obtain this species on Auyan-tepui is perplexing. Nevertheless, our 
establishment of the presence of the Tepui Tinamou there and its 
presumptive presence on Chimanta suggest that this species has a much 
wider distribution in the eastern Pantepui than had been previously 
recognised. 


TURKEY VULTURE Cathartes aura 
Individuals frequently were seen soaring near the tepui rim. 


BLACK VULTURE Coragyps atratus 
One sight record at Angel Falls (Camp IV). 


ROADSIDE HAWK Buteo magnirostris 
This hawk was seen at several localities at elevations from 1700 to 


2200 m. 


GREAT BLACK HAWK Buteogallus urubitinga 


One individual was seen soaring out of the tepui’s central canyon and 
over a forested area of the tepui at 1700 m. 


SWALLOW-TAILED KITE Elanoides forficatus 


Up to three individuals were observed, both flying and perched; in 
the vicinity of Angel Falls. 


G. F. Barrowclough et al. 196 Bull. B.O.C. 1997 117(3) 


LITTLE CHACHALACA Ortalis motmot 
Several individuals were seen and heard calling at 1530 m. 


SPOTTED SANDPIPER Actitis macularia 


This North American migrant was observed foraging along a large 
river near the rim of the central canyon. 


RED AND GREEN MACAW Ara chloroptera 
Daily sight records of a pair at 1530 m (near Camp ITI). 


BUFF-FRONTED OWL Aegolius harrisi 


A male with small testes and light body moult was collected at night 
in a mistnet in an open rocky area at 1700 m. This little known owl had 
been recorded in Venezuela from a few widely scattered localities: 
Mérida in the Andes, El Junquito in the Distrito: Federal (unpubl. 
record; specimen at AMNH), and in the Pantepui from Cerro de la 
Neblina (Willard et al. 1991). This is the easternmost record of the 
nominate northern race. This record and the recent one from Neblina 
indicate that intensive nocturnal use of mistnets may establish a general 
occurrence of this species on the larger, higher tepuis. 


OILBIRD Steatornis caripensis 


One specimen of this wide-ranging bird was taken by mistnet over a 
stream at 1750 m where 15 to 20 birds were attracted to a light used for 
collecting moths on a foggy night. The species is known from scattered 
localities throughout the Pantepui (Willard et al. 1991) as well as 
elsewhere in South America. 


TEPUI SWIFT C'ypseloides phelpsi 

Although Auyan-tepui is the type locality for this species (Collins 
1972), it had not been previously recorded on the summit plateau. 
Gilliard (1941) reported it to be very common on the slopes up to 
1100 m with flocks of up to 5000. On the summit it commonly was seen 
foraging low over vegetation, streams, and open rock. ‘wo females and 
three males were collected; all had moderately enlarged gonads and no 
moult. 


WHITE-COLLARED SWIFT Streptoprocne zonaris 


This widely distributed swift frequently was seen in large flocks of 
up to several hundred birds flying high over the tepui. 


BROWN VIOLETEAR Colibri delphinae 

This species has an altitudinal range from tropical to temperate zones 
and a wide distribution in northern South America; it is known from 
many tepuis (Willard et al. 1991). This hummingbird was uncommon, 
with only two specimens taken by mistnet. Neither specimen was in 
moult; gonads were small. 


VELVET-BROWED BRILLIANT Heliodoxa xanthogonys 

This widespread Pantepui endemic was uncommon; two specimens 
were taken by mistnet. A male had moderately enlarged testes; a female 
had minute ovaries; both were in body moult. 


G. F. Barrowclough et al. 197 Bull. B.O.C. 1997 117(3) 


VERSICOLORED EMERALD Amazilia versicolor 


Two specimens were collected by mistnet at 1700m. Neither 
specimen had enlarged gonads; a male was moulting tail feathers and a 
female had no moult. 


COPPER-RUMPED HUMMINGBIRD Amazilia tobaci 


One specimen (2) was collected on the summit at 1700 m. It was 
previously known from the southern slopes of this tepui up to 1500 m. 
The bird had small ovaries and was in body moult. 


RUFOUS-TAILED TYRANT Knipolegus poecilurus 


This flycatcher is widespread in the Pantepui. A specimen collected 
at c. 2400 m on the south rim by an expedition from the Universidad 
Central de Venezuela in 1956, deposited at EBRG, is the only record of 
this species from Auyan-tepuli. 


LESSER ELAENIA Elaenia chiriquensis 


Two male specimens of this widespread species were collected at 
1700 and 2200 m; neither had enlarged gonads or moult, but both were 
rather fat. It was previously known from the slopes up to 1100m 


(Gilliard 1941). 


BLACK-BILLED THRUSH Turdus ignobilis 


This thrush was fairly common in wooded areas. Three of five males 
had enlarged testes and one of two females had a moderately enlarged 
ovary. None of the birds was in moult. The Phelps Venezuela 
Expedition collected this species on the south slopes up to 1500m 
(Gilliard 1941), but did not find it on the summit. 


RED-SHOULDERED TANAGER Tachyphonus phoenicius 


Gilliard (1941) collected this species from 460 m on the savanna to 
1800 m on the slopes. We took an adult male with moderately enlarged 
gonads at 1700 m on the plateau. 


Acknowledgements 


The authors are grateful to Armando Michelangeli Ayala, President of Fundacion 
Terramar, for organisation and logistic support. The Venezuelan Air Force provided 
essential flights of personnel and equipment. Gilberto Pérez Ch. and Gregorio Lozano 
assisted in the collection and preparation of specimens in the field. Fellow expedition 
members Petia Alcécer, Anibal Chacon, John W. Daly, Maureen A. Donnelly, José Luis 
Garcia, Ricardo Guerrero, Fabian Armando Michelangeli H., Charles W. Myers, and 
Adriana Sanchez H. assisted our work in many ways and helped to create efficient and 
pleasant field camps. Luis Pérez Ch. assisted in our examination of specimens at the 
Coleccion Ornitologica Phelps. The staff of the Colecciones de la Estacién Biolégica de 
‘Rancho Grande enabled one of us (M.L.R.) to examine specimens from the 1956 
expedition. Principal funding was provided by Robert G. Goelet, with additional support 
from the L. C. Sanford Trust, the L. J. Sanford Trust, and the Phelps Foundation. 
Permits for field work were provided by INPARQUES and PROFAUNA. Charles W. 
Myers provided comments on parts of the manuscript and Sra Kathy D. de Phelps 
graciously assisted us in many ways. 


G. F. Barrowclough et al. 198 Bull. B.O.C. 1997 117(3) 


References: 

Barrowclough, G. F., Escalante-Pliego, P., Aveledo-Hostos, R. & Pérez-Chinchilla, L. 
1995. An annotated list of the birds of the Cerro Tamacuari region, Serrania de 
Tapirapeco, Federal Territory of Amazonas, Venezuela. Bull. Brit. Orn. Cl. 115: 
211-219. 

Berry, P. E., Holst, B. K. & Yatskievych, K. 1995. Flora of the Venezuelan Guayana. 
Vol. 1: Introduction. 'Timber Press, Portland, Oregon. 

Chapman, F. M. 1939. The upper zonal birds of Mt. Auyan-tepui, Venezuela. Amer. 
Mus. Novit. 1051. 

Collins, C. T. 1972. A new species of swift of the genus Cypseloides from northeastern 
South America (Aves: Apodidae). Contr. Sci. Nat. Hist. Mus. Los Angeles Co. 229. 

Gilliard, E. T. 1941. The birds of Mt. Auyan-tepui, Venezuela. Bull. Amer. Mus. Nat. 
Hist. 77: 439-508. 

Mayr, E. & Phelps, W. H., Jr 1967. The origin of the bird fauna of the south Venezuela 
highlands. Bull. Amer. Mus. Nat. Hist. 136: 269-328. 

Medina Cuervo, G. 1992. La avifauna del macizo del Chimanta. Pp. 281-294 in O. 
Huber (ed.), El Macizo de Chimanta. Oscar Todtmann Editores, Caracas. 

Meyer de Schauensee, R. & Phelps, W. H., Jr 1978. A Guide to the Birds of Venezuela. 
Princeton Univ. Press. 

Phelps, W. H. & Phelps. W. H., Jr 1958. Lista de las aves de Venezuela con su 
distribucion. No passeriformes. Bol. Soc. Venez. Cienc. Nat. 90: 1-317. 

Steyermark, J. A. 1967. Flora del Auyan-tepui. Acta Bot. Venez. 2: 5-370. 

Tate, G. H. H. 1938. Auyantepui. Notes on the Phelps Venezuela expedition. Geogr. 
Rev. 28: 452-474. 

Willard, D. E. et al. 1991. The birds of Cerro de la Neblina, Territorio Federal 
Amazonas, Venezuela. Fieldiana 65: 1-80. 


Addresses: George F. Barrowclough & Paul R. Sweet, Department of Ornithology, 
American Museum of Natural History, Central Park West at 79th Street, New York, 
NY 10024, U.S.A. Miguel Lentino R., Coleccién Ornitologica Phelps, Apartado 
2009, Caracas, Venezuela. 


© British Ornithologists’ Club 1997 


Appendix 1 


Coordinates and elevations of 1994 collecting localities. 


Camp I. 5°51'N, 62°32’W, 1700 m. 
Camp II. 5°54'N, 62°29’W, 1750 m. 
Camp ITI. 5°53’N, 62°38'W, 1850 m. 
Camp IV. 5°58’N, 62°33’W, 1700 m. 
Camp V. 5°46'N, 62°32'W, 2200 m. 


Appendix 2 


Complete list of birds known from the summit plateau of Auyan-tepui, Estado Bolivar, 
Venezuela. *=sight record. 


Crypturellus ptaritepui, Podiceps dominicus, Cathartes aura*, Coragyps atratus*, Buteo 
magnirostris*, Buteogallus urubitinga*, Elanoides forficatus*, Ortalis motmot*, Actitis 
macularia*, Gallinago paraguaiae*, Columba fasciata, Ara chloroptera*, Nannopsittaca 
panychlora, Aegolius harrisu, Steatornis caripensis, Caprimulgus longirostris, Streptoprocne 
zonaris*, Cypseloides phelpsi, Aeronautes montivagus*, Campylopterus hyperythrus, Colibri 
delphinae, Colibri coruscans, Polytmus milleri, Amazilia versicolor, Amazilia tobaci, 
Heliodoxa xanthogonys, Piculus rubiginosus, Cranioleuca demissa, Premnoplex adusta, 
Automolus roraimae, Lochmias nematura, Thamnophilus insignis, Myrmothera simplex, 
Chloropipo uniformis, Knipolegus poecilurus, Hirundinea ferruginea*, Todirostrum 
russatum, Mecocerculus leucophrys, Elaenia chiriquensis, Elaenia dayi, Elaenia pallatangae, 
Troglodytes rufulus, Turdus olivater, Turdus ignobilis, Macroagelaius imthurm, Myioborus 
castaneocapillus, Coereba flaveola, Diglossa major, Tangara cyanoptera, Tachyphonus 
phoenicius, Catamenia homochroa, Atlapetes personatus, Zonotrichia capensis. 


C. B. Frith & D. W. Frith 199 Bull. B.O.C. 1997 117(3) 


A distinctive new subspecies of Macgregor’s 
Bowerbird (Ptilonorhynchidae) of 
New Guinea 


by Clifford B. Frith S& Dawn W. Frith 


Received 30 March 1996 


Macgregor’s Bowerbird Amblyornis macgregoriae is the most widely 
distributed of the four Amblyornis species of ‘maypole’ bower-building 
‘gardener’ bowerbirds, all confined to mainland New Guinea. 
Macgregor’s Bowerbird is found throughout the main mountain ranges 
of New Guinea except those of the extreme west of Irian Jaya (Tamrau, 
Arfak and Wandammen Mts.) at altitudes of 1600-3300 (rarely 
1200-3300) m (Gilliard 1969, Schodde & McKean 1973, Beehler et al. 
1986). Adult males are adorned with an extensive bright orange crest 
which they display to females during courtship at bowers they 
construct of sticks placed about a central sapling and decorated with 
moss and various objects including fruits, flowers and leaves (Frith 
1970, Cooper & Forshaw 1977, Borgia 1986). 

In the course of a study of bowerbird specimens in several major 
museum collections about the world a series of eight skins of 
Macgregor’s Bowerbird from Mt. Bosavi, Southern Highlands, Papua 
New Guinea, were examined at the Bernice Pauahi Bishop Museum 
(BPBM), Honolulu, Hawaii, in January 1995. These consisted of four 
adult (fully crested) males, three immature (uncrested) males and one 
adult female, and were all collected during 6—9 June 1973 at c. 1400 m 
on the NNW slopes of Mt. Bosavi in forest. Seven were recorded by 
the preparator as caught in “‘snap traps’’ set to catch live mammals and 
one was mist-netted by A. B. Mirza. 

We again examined all eight BPBM specimens from Mt. Bosavi at 
the American Museum of Natural History (AMNH), New York, in 
February 1995 where direct comparisons could be made with 
specimens from the nearest collection locality for the species, the Mt. 
Hagen area, Eastern Highlands, Papua New Guinea. This AMNH 
material, of the subspecies A. m. kombok (Schodde & McKean 1973), 
was extensive enough (see below) to permit meaningful conclusions 
concerning any differences in morphology between the birds of the Mt. 
Bosavi and Mt. Hagen area samples. For comparative purposes the 
colours of Smithe (1975) are referred to by their capitalized names with 
the numbers allocated to them in parenthesis. Where we deem a colour 
~name of Smithe (1975) inappropriate we do not capitalise the name 
we use. 

All wing and tail measurements were made with the same rulers, all 
others with the same electrical digital calipers, and all by CBF. Wing 
length measured was the flattened and straightened, thus maximised, 
chord, using a stopped steel rule. Tail length was measured from the 
point of entry of the central pair into the bird’s skin to the tip of the 
longest feather with a small steel rule. Bill length was measured from 


C. B. Fnth & D. W. Frith 200 Bull. B.O.C. 1997 117(3) 


ePaiyer 
River 


Hagen 


Giluwe K 
a eGoroka 


Lake 
Kutubu, eAwande 
Bosavi Karimui 


Figure 1. Map showing the location of Mt. Bosavi, type locality of Amblyornis 
macgregoriae lecroyae, and other locations in Papua New Guinea mentioned in the text. 
Solid triangles, mountains; solid dots, places. After Diamond (1972). 


the bill tip to the cranio-facial hinge, and bill width was taken at the 
anterior edge of the nostril. Tarsus length was that of the 
tarsometatarsal bone. 

Macgregor’s Bowerbird was previously known to occur on Mt. 
Bosavi as Bell (1975) reported seeing it there in August 1970 and Thane 
Pratt (unpubl. data zn litt.) also saw the species there in February 1975, 
but it appeared to have remained uncollected there until the BPBM 
specimens detailed here were taken. We were much surprised, 
therefore, to discover in June 1995 a single adult male specimen 
collected on 30 August 1970 at 1525 m a.s.l. on Mt. Bosavi by H. L. 
Bell within the collections of the Papua New Guinea National Museum 
and Art Gallery (PNGM), Port Moresby. Bell had prophetically 
written in pencil on his field specimen label ‘“‘new subspecies?”’. 

Mt. Bosavi is a 2600 m mountain located in the Western District of 
Papua New Guinea north of the Fly River delta of southern Papua New 
Guinea (Fig. 1), isolated from the central mountains of New Guinea as 
a vast discrete extinct volcanic cone in extensive lowlands. The nearest 
known population of Macgregor’s Bowerbird to that on Mt. Bosavi is 
on the western side of the Tari Valley (Frith & Frith 1992) where a 
single specimen was collected for the BPBM and another for the 
Australian Museum (AM). Whilst we had previously examined and 
measured these two Tari Valley specimens they were not available for 
direct comparative study with the Bosavi series. In any event, little 
could be concluded from such limited material. 

The generally brownish plumage of all eight BPBM birds from 
Mt. Bosavi is consistently and strikingly more saturated with reddish 
pigment (136) to give them an overall different appearance to a series of 


C. B. Frith & D. W. Frith 201 Bull. B.O.C. 1997 117(3) 


A.m. kombok birds consisting of the following: 13 (8 adult ¢d, 2 
immature gd, 2 29, 1 unsexed) from Mt. Hagen at c. 8500 feet, three 
(1 adult ¢, 1 immature ¢, 1 9) from Nondugl, two (1 adult 4, 
1 immature 4) from Mt. Karimui, one adult ¢ from Awande and four 
(3 adult $4, 1 immature ¢) from Kubor Mts. (see Fig. 1 for locations). 
The strikingly more reddish plumage of the Bosavi birds is clearly 
indicative of differentiation and is not a product of any kind of 
post-mortem colour changes described and discussed by Gilliard & 
LeCroy (1961). The Mt. Bosavi birds are, sex for sex and age group for 
age group, consistently a little smaller in both wing and tail length than 
A.m. kombok (Table 1). Gilliard & LeCroy (1961) stressed the use of 
tail (and crest) length to differentiate subspecies of A. macgregoriae. 
Adult male crest length measured from posterior base (Gilliard & 
LeCroy 1961) does not differ between Bosavi birds and A. m. kombok, 
but in exposed crest length the Bosavi birds may show a smaller area of 
colour. The latter measurement is less objective than the former, being 
a measure of the length of colourful crest viewed from above, but it is 
possible that Bosavi birds have more dark fore-crown feathering 
resulting in a shorter visible crest (Table 1). 

As we agree with Gilliard (1969), Diamond (1972) and Schodde & 
McKean (1973) that the birds from the Kubor Mts., Mt. Karimui, 
Awande, Nondugl and Mt. Hagen are all clearly attributable to the 
subspecies A. m. kombok, we confine our comparisons of the plumage of 
the Mt. Bosavi birds to that of the 16 from Mt. Hagen and Nondugl. A 
general description of these 16 specimens follows: 


Underparts somewhat variable in degree of colour saturation (worn birds being 
slightly to considerably paler) but typically rich Buff (24) to Clay Color (26), and to 
Cinnamon (123A) in only two specimens. An indistinct broad area of the chest to the 
upper breast somewhat darker and browner, close to Raw Umber (23), but variable. 
Throat slightly paler than abdomen. Underwing coverts pure Clay Color (123B) and 
trailing edge of the primaries a more dilute tone of this colour. Forecrown, mantle and 
back of adult males Raw Umber (23) to Prout’s Brown (121A) and crown of immature 
males and females redder, like Amber (36) but darker. Wings and upper tail less rich, 
more an Olive-Brown (28). Crest colour of the adult males variable, the richest being 
Spectrum Orange (17) in specimen 705724 and others paler, possibly to some degree 
due to fading. 


Comparative photographs, taken under similar conditions by us, of the 
eight Mt. Bosavi specimens at the BPBM and of the single specimen at 
the PNGM enable us to determine that the latter is of simular 
appearance to the former. Notwithstanding the prior collection date of 
Bell’s single specimen we designate it as a paratype because the BPBM 
series of eight clearly represents a far sounder basis for formal 
description of the holotype. A comparative description of the Mt. 
Bosavi population of A. macgregoriae follows: 


Amblyornis macgregoriae lecroyae subsp. nov. 


Holotype. Bernice Pauahi Bishop Museum, Honolulu, Hawaii, 
BBM-NG 103303, adult male, from NNW slopes of Mt. Bosavi, 
Southern Highlands Province, Papua New Guinea, 1400 +m, collected 


C. B. Fnth & D. W. Frith 202 Bull. B.O.C. 1997 117(3) 


TABLE 1 
Measurements of all known specimens of Amblyornis macgregoriae lecroyae and of sexed 
specimens of A. m. kombok in several museum collections 


Exposed 
Wing Tail Tarsus Bill Bill Crest crest 
length length length length width length? length* 
A. m. lecroyae' 
Adult males 
BBM-NG 103303 Type 131 82 37.0 28.5 8.5 63 75 
BBM-NG 103327 130 78 34.8 28.5 re 63 74 
BBM-NG 103328 135 84 36.4 25.9 fide) 66 67 
BBM-NG 103329 132 84 333 Dead, Paik 65 83 
PNGM B937/25303 132 80 37.4 28.7 8.8 62 83 


Mean, s.d. 132, 1.87 ° 82,°2.58 ° 36:8, 0.95 27-9) °T.16" (8.0,°0.57_ ‘GHGs Gt O.b+ 


Immature males 


BBM-NG 103330 131 82 34.0 204 7.1 
BBM-NG 103331 133 83 — 28.3 4:9 
BBM-NG 103337 129 80 38.9 2 Tard had 
Mean, s.d. 131, 2.00 82, 1.20 36.5, 3:46 27.6, 0.59 - 7.4, 0.42 


Adult female 
BBM-NG 103339 129 81 37.9 26.6 8.1 


A. m. kombok? 
Adult males 


Mean, s.d. 137, 3.92 87, 3.30 37.4, 2.03 28.1,0.85 7.4,0.55 63, 4.98 84, 5.59 
n 32 29 32 32 31 32 
Immature males 

Mean, s.d. 135.3566) 85.280) 37.8.0. 9745 27:8. 0429). 75 uLOU 

n 17 Wy) 17 17 17 

Adult females 

Mean, s.d. 131, 3.03 84, 2.91 36.2,1.45 28.7, 0.73 8.6, 0.62 

n 11 11 11 11 11 


'BBM-NG=Bishop Museum, Honolulu; PNGM=Papua New Guinea National Museum and Art 
Gallery, Port Moresby. 

The 60 specimens of A. m. kombok include: 25 from American Museum of Natural History, New 
York; 12 Natural History Museum, Tring; 10 Australian National Wildlife Collection, CSIRO, 
Canberra; 6 Australian Museum, Sydney; 5 PNGM; 1 BPBM;; and 1 Field Museum of Natural 
History, Chicago. 

3From posterior base (cf. Gilliard & LeCroy 1961, Schodde & McKean 1973). 

*The length of visible crest colour viewed and measured from above. 


by ‘pan’ trap by A. B. Mirza on 6 June 1973. See Table 1 for 
measurements. 

Diagnosis. Upperparts: forecrown, side of face, malar, crown and 
mantle much richer and more strongly suffused with orange than in 
A.m. kombok (see above), being closest to Raw Umber (23) but slightly 
darker and suffused with rich brown-orange. Wings and tail less olive 
and more brown-orange than in A. m. kombok, the former being closer 


C. B. Frith & D. W. Frith 203 Bull. B.O.C. 1997 117(3) 


to but darker than Cinnamon-Brown (33) and the latter closer to Raw 
Umber (23). Crest colour as the most heavily saturated specimen of 
A.m.kombok, being Spectrum Orange (17). Underparts: conspicuously 
more orange-brown than all above specimens of A. m. kombok, and 
particularly so on the chin, throat and the darker area of the chest and 
upper breast. Throat closest to Raw Umber (123) and the darker 
chest/breast area, sides of abdomen and the flanks closest to Cinnamon 
Brown (33) but washed orange; these darker areas contrasting slightly 
with the paler Cinnamon (123A) remainder of the underparts. 
Underwing coverts pale rich ginger (as Clay Color 123B, but much 
more orange) and trailing edge of the primaries a more dilute tone of 
this colour. 

Paratypes. Adult male specimens BBM-NG 103327, 103329 and 
PNGM B937/25303 are very similar to the holotype specimen but 
BBM-NG 103329 has a deeper orange crest. Specimen BBM-NG 
103328 is even more orange, almost red-brown above, and its crest is as 
103320. All three immature males and the female Mt. Bosavi birds 
(Table 1) are plumaged as the adult males but (lacking a crest) have the 
forecrown and crown slightly darker and more richly suffused orange 
than the rest of the upperparts and contrast with them slightly. 

Distribution and habitat. Presently known only from the forest of the 
type locality and at altitudes of c. 1400 and 1525 maz.s.l. While this new 
subspecies is presently known only from Mt. Bosavi the use of this 
geographical feature as a trinomial is not desirable or appropriate as 
similar birds may prove to be found elsewhere. The southern end of the 
Muller Range is a possibility, though an unlikely one. 

Comments. A. m. lecroyae is distinctly different in overall appearance 
to A. m. kombok in being darker and significantly more brown-orange. 
Of the above-described A. m. kombok series only female-plumaged 
(crestless) specimen AMNH 705730 from Mt. Hagen of uncertain sex 
approaches the coloration of the Mt. Bosavi A. m. lecroyae series, being 
similar in colour and tone above except for the upper tail which is not 
as dark or orange. Its crown is similar to but perhaps not quite as dark 
and orange as in female-plumaged A. m. lecroyae. Its underparts are 
generally similar to A. m. lecroyae but the throat is paler and the 
remainder, including the underwing, not as dark or as orange-suffused 
as the latter subspecies. Thus adult male, immature male and female 
A.m. lecroyae are conspicuously more orange-suffused and darker in 
their plumage than A. m. kombok. T. Pratt (in litt.) handled one mist- 
netted live adult male Macgregor’s Bowerbird on Mt. Bosavi and wrote 
of it at the time “‘had a very short crest—molting?’’, which is noteworthy 
in view of his considerable experience with the species elsewhere. 

We have found no description of the form of bower built by male 
Macgregor’s Bowerbirds on Mt. Bosavi, but as Bell (1975) and T. Pratt 
(zn litt.) made no remark in this regard and as bowers do not appear to 
differ in general appearance between the other populations of the 
species, we suspect Mt. Bosavi bowers are not significantly different in 
general appearance. We would not hesitate to elevate the Bosavi birds 
to full species status, however, should they prove to build a consistently 
different bower to those of other populations of the species, given the 


C. B. Fnth & D. W. Frith 204 Bull. B.O.C. 1997 117(3) 


species-specific nature of bowerbirds’ bowers (Diamond 1982, Borgia 
1986). 

Etymology. It is with great pleasure that we name this distinctive 
orange form of Macgregor’s Bowerbird for our colleague and friend 
Mary LeCroy of the Department of Ornithology, American Museum of 
Natural History, New York. This is by way of small acknowledgement 
of her significant contributions to the ornithology of New Guinea and 
elsewhere, and much generous help long given to numerous 
ornithologists working on bowerbirds and other birds of New Guinea, 
and elsewhere. 


In addition to raising the possibility that his single specimen of 
Macgregor’s Bowerbird from Mt. Bosavi represented a new subspecies 
(see above), H. L. Bell (1975) noted that he took samples of three small 
passerine species from there large enough to indicate to him that, “if 
not distinct sub-species, they are definitely of different size to ‘typical’ 
birds on the main range’’. Moreover, Schodde & Weatherly (1983) 
described a new species of fairy-wren Malurus campbelli, subsequently 
considered a distinctive subspecies of M. grayi by LeCroy & Diamond 
(1995), discovered at c. 700-800m a.s.l. on Mt. Bosavi. These 
observations, taken together with the distinctive morphology of 
Amblyornis macgregoriae lecroyae on Mt. Bosavi, suggest the possibility 
of other undescribed avian taxa awaiting discovery on that mountain. 


Acknowledgements 


For kindly making available collections in their care and for facilities to study them in situ 
or elsewhere we sincerely thank Allen Allison and Carla Kishinami of the Bernice Pauahi 
Bishop Museum, Honolulu; the Chapman Fund, Dean Amadon, Chris Blake, Walter 
Bock, Joel Cracraft, Mary LeCroy, Manny Levine, Lester Short and Richard Sloss of 
the American Museum of Natural History, New York; Richard Schodde, Ian Mason 
and John Wombey, Australian National Wildlife Collection, Canberra; Walter Boles and 
Wayne Longmore, Australian Museum, Sydney; Frank Bonaccorso, Ilaiah Bigilale and 
Paul Wanga, National Museum and Art Gallery of Papua New Guinea, Port Moresby; 
Robert Prys-Jones, Peter Colston and Michael Walters, Natural History Museum, 
Tring; David Willard, Field Museum of Natural History, Chicago. Allen Allison, Carla 
Kishinami, Mary LeCroy, Thane Pratt and Richard Schodde kindly provided 
constructive thoughts and comments on a draft of this contribution. 


References: 

Beehler, B. M., Pratt, T. K. & Zimmerman, D. A. 1986. Birds of New Guinea. Princeton 
Univ. Press. 

Bell, H. L. 1975. Mount Bosavi as an ecological island. New Guinea Bird Society 
Newsletter 110: 8-10. 

Borgia, G. 1986. Sexual selection in bowerbirds. Scientific American 254: 92-98. 

Cooper, W. T. & Forshaw, J. M. 1977. The Birds of Paradise and Bower Birds. Collins, 
Sydney. 

Diamond, J. M. 1972. Avifauna of the Eastern Highlands of New Guinea. Publ. Nuttall 
Orn. Club No. 12. 

Diamond, J. M. 1982. Evolution of bowerbirds’ bowers: animal origins of the aesthetic 
sense. Nature 297: 99-102. 

Frith, C. B. 1970. Sympatry of Amblyornis subalaris and A. macgregoriae in New Guinea. 
Emu 70: 196-197. 

Frith, C. B. & Frith, D. W. 1992. Annotated list of birds in western Tari Gap, Southern 
Highlands, Papua New Guinea, with some nidification notes. Australian Bird 
Watcher 14: 262-276. 


S.A. Lousada & S. N. G. Howell 205 Bull. B.O.C. 1997 117(3) 


Gilliard, E. T. 1969. Birds of Paradise and Bower Birds. Weidenfeld & Nicolson, London. 

Gilliard, E. T. & LeCroy, M. 1961. Birds of the Victor Emanuel and Hindenburg 
Mountains, New Guinea. Results of the American Museum of Natural History 
Expedition to New Guinea in 1954. Bull. Amer. Mus. Nat. Hist. 123: 1-86. 

LeCroy, M. & Diamond, J. 1995. PLumage variation in the Broad-billed Fairy-wren 
Malurus grayi. Emu 95: 185-193. 

Schodde, R. & McKean, J. L. 1973. Distribution, taxonomy and evolution of the 
gardener bowerbirds Amblyornis spp. in eastern New Guinea with descriptions of 
two new subspecies. Emu 73: 51-60. 

Schodde, R. & Weatherly, R. G. 1983. Campbell’s fairy-wren Malurus campbelli, a new 
species from New Guinea. Emu 82: 308-309. 

Smithe, F. B. 1975. Naturalist’s Color Guide. American Museum of Natural History. 


Address: C. B. & D. W. Frith, “‘Prionodura’’, P. O. Box 581, Malanda, Queensland, 
Australia 4885. 


© British Ornithologists’ Club 1997 


Amazona oratrix hondurensis: a new subspecies 
of parrot from the Sula Valley of northern 
Honduras 


by Sebastian A. Lousada & Steve N. G. Howell 
Received 31 May 1996 


Although Monroe & Howell (1966) suggested that there was a new 
subspecies of yellow-crowned parrot (i.e. a member of the Amazona 
ochrocephala complex; see General remarks) in the Sula Valley of 
Honduras, they had insufficient data to formally describe it at that time. 
Following our field and museum work, the complex situation in 
northern Honduras has been clarified. Most of the details of this were 
_described by Lousada & Howell (1996) and, despite few museum 
specimens, we believe there is now ample evidence to describe the 
distinctive dimorphic Sula Valley birds as a new subspecies: 


Amazona oratrix hondurensis, subsp. nov. 


Holotype. Museum of Comparative Zoology, No. 136569; female. 
Honduras: Depto. Cortes; c. 12 miles NE of La Lima, along the 'Toloa 
canal at Urraco; 10 March 1928; collected by E. Bangs, original field 
number J. L. Peters 5505. 

Diagnosis (adults). Although clearly a member of the Amazona 
ochrocephala complex, hondurensis differs from belizensis and 
“guatemalensis”’ (an undescribed dimorphic form of belizensis; Lousada 
& Howell 1996) in having yellow head coloration limited to the 
forehead and crown only, and in frequently having a yellow nape patch. 
Differs from caribaea in its overall paler bill and more extensive yellow 
on the forehead and crown. This latter is usually in a rounded or 
triangular shape versus a narrower band in caribaea. Caribaea has dark 
cere/bristle coloration, while hondurensis is often far paler due to the 


S.A. Lousada & S. N. G. Howell 206 Bull. B.O.C. 1997 117(3) 


presence of at least some blond bristles and paler underlying skin. 
Differs from parvipes in its consistently pale bill and extensive yellow 
forehead and crown (see Figure 2c, page 47, Lousada & Howell 1996). 
The yellow-crowned morph of hondurensis is similar to panamensis from 
which it differs in its larger size and paler bill (banamensis usually has 
darker coloration on the bill). 

Measurements (mm) of holotype. Wing (chord) 212, tail 100, culmen 
(from nostril) 30, length of middle toe (without nail) 30. 

Distribution. Originally ranged throughout the Sula Valley of 
northern Honduras and west and east along the Caribbean coastal 
lowlands. It is now apparently restricted to a small area west of Tela 
and south of Punta Sal in the lower Sula Valley. | 

Etymology. We name this subspecies hondurensis recognising that it is 
the only member of the A. ochrocephala complex endemic to mainland 
Honduras. 

Specimens examined. Honduras: Chasniguas (CMNH 1 male); Urraco 
(MCZ 1 female); Laguna 'Toloa (ANSP 1 male); 3mi E La Lima 
(LSUMZ 1 female). 

Captive specimens examined. Generally, captive birds are not included 
aS specimens examined in a taxonomic description. In this case, 
however, we feel that they can add valuable data due to the limited 
series of museum skins, the undesirability of collecting more specimens 
from a very small remnant population, and the fact that some collection 
data are available for the birds noted below. The problem with captive 
birds is that in most cases the exact or even general location of 
collection from the wild is unknown, especially in the case of parrots in 
the international trade. Nevertheless, our experience in this region 
indicates that internal trade, while very common, generally reflects 
birds available locally from the wild. This is particularly true in the 
case of poorer campesinos (country people). Amongst these households 
it is very rare to find pet birds that are any significant distance away 
from their natural ranges. 

In all, 17 wild-caught captive birds (4 males, 7 females, 6 unknown) 
have been studied. Six of these were seen in Honduras, within or 
bordering the present range of hondurensis. 'The remaining nine birds 
were seen in the United States. Fortunately, the importer is known for 
these birds and he reported collecting them from “the Puerto Cortes 
area around 1980” (J. McDonald pers. comm.), an area bordering the 
bird’s present range but well within the range at that time. 

Field observations. As published previously (Lousada & Howell 1996), 
Lousada and Morales located a flock of approximately 75 hondurensis 
in the wild. Although field observations are necessarily subjective, they 
took photographs which, while not of high quality, do show the key 
features of yellow crowns and pale bills. 

Other specimens of the complex examined. Although many other 
specimens of this complex have been examined previously, the 
following specimens were all brought together to be examined side by 
side specifically for this paper. 

oratrix. Mexico: Rio Cruz, Tampa (MCZ 1 female); Pasa Nueva, 
Veracruz (MCZ 1 male, 1 female); Petapa, Tehuantepec (MCZ 1 male). 


S.A. Lousada & S.N. G. Howell 207 Bull. B.O.C. 1997 117(3) 


belizensis. Belize: Gallon Jug, Orange Walk Dist. (LSUMZ 1 male); 
Hill Bank, Orange Walk Dist. (LSUMZ 1 female); near Ycacos Lagoon 
(MCZ 1 male); All Pines (CMNH 1 male). 

caribaea. Honduras: Isla Barbareta (CMNH 2 males, 1 female); Isla 
Guanaja (CMNH 1 male). 

parvipes. Nicaragua: 25 mi. SE Leicus Creek (UCLA 1 male); Leicus 
Creek (UCLA 3 females); 23 km SSW Waspam (UCLA 1 male, 1 
female); 15 km SSW Waspam (UCLA 1 male, 1 female); 56 mi NW 
Puerto Cabeza (UCLA 1 female). 

auropalliata. Mexico: 6 mi. NE Tonala, Chiapas (LSUMZ 1 female); 
Guatemala: 10 mi. SW Taxisco, Santa Rosa (LSUMZ 2 females); 
Finca el Cipres (MCZ 1 female); “‘“Guatemala’ (ANSP 2 unsexed); 
Honduras. San Lorenzo (MCZ 1 female); 7 mi. NE~ Choluteca 
(LSUMZ 1 male); Nicaragua: Coseguina (ANSP 1 _ female); 
“Nicaragua” (MCZ 1 male); Costa Rica: Lenorio (MCZ 1 male, 2 
females); ““Costa Rica’? (MCZ 1 male, 1 female). 

panamensis. Panama: Divala (MCZ 1 male); Parita (CMNH 1 male); 
unspecified location (MCZ 1 unsexed). 


Individual variation 

Individual variation is a marked feature of this population, as it is of 
some other Amazona taxa. In what follows, capitalized colour 
descriptions and numbers follow Smithe (1975, 1981), although almost 
none of them matches exactly. Apart from some key areas of colour on 
the head, overall colours are fairly typical for the complex. Individual 
variation of the subspecies as a whole has been published previously 
(Lousada & Howell 1996). The following are notes primarily on the 
museum specimens. The forehead and crown of the holotype have a 
pale yellow patch, nearest Sulfur Yellow (157), 40 mm in length from 
the cere and 20 mm wide at its broadest point with a total area of 
492 mm”. The crown coloration of the other three specimens is similar 
but is more extensive (532, 796 and 916 mm?” in ANSP 90568, LSUMZ 
29066 and CMNH 20448 respectively) and in a more rounded shape. 
(ANSP 90568 is missing many crown feathers and originally may well 
have had more yellow than measured). ANSP 90568 and LSUMZ 
29066 lack yellow nape patches and may be immatures, or more mature 
individuals of the yellow-crowned morph. The holotype has a spotty 
yellow nape patch on the nape and hindneck which is darker yellow 
than the crown, closest to Spectrum Yellow (55). CMNH 20448 has a 
more sizeable yellow nape patch than the holotype; due to yellow flecks 
on the occiput this almost joins the crown patch. (One of the captive 
specimens shows this to an even more exaggerated extent.) Dusky 
margins on the feathers vary from bird to bird (as in other subspecies of 
the complex), being especially conspicuous in LSUMZ 29066. 
Immatures often have more dusky margins than older birds but this is 
not consistent enough to be a reliable age indicator. Although the 
thighs on all four specimens are green, there are some yellow flecks at 
the base of the thighs of CMNH 20448. The amount of red and yellow 
on the bend of the wing also varies among the specimens. There is none 
obvious on the bend of the wing of ANSP 90568 (another possible sign 


S.A. Lousada & S. N. G. Howell 208 Bull. B.O.C. 1997 117(3) 


TABLE 1 
Area of yellow in crown (in mm?) of adults of some subspecies of the Amazona 
ochrocephala complex 


Subspecies n Range Mean 
belizensis 4 2264-2546 2460.5 
hondurensis 4 492-916 684 
caribaea 3 304-502 426 
parvipes 8 0-514 160.25 | 
auropalliata (all specimens examined) 14 0-398 128.36 
auropalliata (Guatemala) 5 35-398 197.8 
auropalliata (Costa Rica) 4 0-142 59 
panamensis - 2 684-748 716 


Notes: Acetate sheets laid over crowns and traced onto. Tracings then laid over 2 mm 
graph paper and counted. Belizensis is the least accurate due to head curvature, and 
includes the eye region. 


of being an immature) and there is only a small amount on the 
holotype: c. 20 mm? (2. feathers) closest to Geranium Pink (13) edged 
with yellow on the right wing and c. 39 mm? (4 red feathers and 1 
yellow) on the left wing. There is more red along the marginal coverts 
that cannot be seen with the wing folded close to the body. The carpal 
edge is green, closest to Lime Green (59). 

It is commonly known that soft-part colours change with death. The 
mandibles of the holotype are closest to Cinnamon (39) and darker at 
the edge and tip of the upper mandible, but were almost certainly far 
paler in life. ‘The mandibles of LSUMZ 29066 are the duskiest, 
perhaps due to bleeding at death and/or the possibility that it may be an 
immature bird. All four specimens have mandibles that are somewhat 
darker than most ovatrix museum specimens. Compared to oratrix 
there may perhaps be some subtle colour differences of one or more of 
the inner layers of the mandibles of hondurensis that are not necessarily 
apparent in life and at least partially account for this. Although the bills 
of the captive hondurensis specimens are all pale, some are a little greyer 
than others. The cere of the holotype has many blond bristles although 
the majority are dark. The ceres of the other specimens have blond 
bristles present in varying quantities. The underlying skin of the ceres 
is also likely to have darkened in death. The captive specimens show 
overall cere colour that ranges from extremely pale (like oratrix) to 
more dusky. 


General remarks 

As mentioned in Lousada & Howell (1996) there is a NW-SE cline 
evident in the amount of yellow in the different allopatric forms of the 
Amazona ochrocephala complex. By measuring crown patterns from 
museum skins, Lousada produced Table 1 which quantifies this. 
Interestingly and recognising the small sample, a similar trend appears 
to occur on the Pacific coast, with southern auropalliata having 
noticeably less yellow on the forehead than northern birds. 


S.A. Lousada & S. N. G. Howell 209 Bull. B.O.C. 1997 117(3) 


There is one other manifestation of the cline that was not mentioned 
in Lousada & Howell (1996). The carpal edge (below the bend of the 
wing) of most Mexican oratrix specimens is brilliant yellow, sometimes 
with a few red and green flecks. This yellow coloration is reduced or 
absent in belizensis. 

Captive “‘guatemalensis’’ showed primarily green to yellowish green 
carpal edges with just occasional yellow or red flecks, as do the rest of 
the specimens of the complex. 

Although AOU (1983) considered Sula Valley birds to be 
Yellow-crowned Parrots A. ochrocephala, we believe that, given a fuller 
understanding of the complex situation in northern Central America 
(Lousada & Howell 1996), hondurensis marks the southern end in a 
cline from Mexican oratrix through belizensis and ‘“‘guatemalensis’’; 
hence, we have placed it with Yellow-headed Parrot (A. oratrix of AOU 
1983). It can even be argued, however, that hondurensis could be placed 
with Yellow-naped Parrot (A. auropalliata of AOU 1983), caribaea 
being intermediate between hondurensis and parvipes (Lousada & 
Howell 1996), or that all of these ‘species’ should be reunited as a 
single, polymorphic species, A. ochrocephala, as treated by Forshaw 


(1973). 


Acknowledgements 


Firstly we thank Raymond A. Paynter (Museum of Comparative Zoology; MCZ) for his 
great kindness in offering to house specimens on loan from other institutions and 
permitting access to the museum’s collection; and the curators and collection managers 
who permitted loans of specimens under their care: Kenneth C. Parkes and Robin Panza 
(Carnegie Museum; CM); Robert S. Ridgely and David Agro (Academy of Natural 
Sciences of Philadelphia; ANSP); J. V. Remsen and S. W. Cardiff (Louisiana State 
University; LSU); Jaap Hillenius (University of California, Los Angeles; UCLA). In 
addition Robin Panza checked some measurements of Carnegie Museum specimens and 
Kenneth C. Parkes reviewed the manuscript and offered many helpful comments. Joe 
McDonald provided information on his importations of hondurensis, Winifred Ewing 
kindly provided a place to stay while working at MCZ, and Isabelle Lousada and Sabra 
Ewing provided logistical support. This is contribution number 712 of the Point Reyes 
Bird Observatory. 


References: 

AOU (American Ornithologists’ Union). 1983. Check-list of North American Birds. 
6th edn. AOU, Washington, D.C. 

Forshaw, J. M. 1973. Parrots of the World. Lansdowne Press, Melbourne, Australia. 

Lousada, S. A. & Howell, S. N. G. 1996. Distribution, variation and conservation of 
Yellow-headed Parrots in northern Central America. Cotinga 5: 46-53. 

Monroe, B. L. & Howell, T. R. 1966. Geographic variation in Middle American parrots 
of the Amazona ochrocephala complex. Occ. Pap. Mus. Zool. Louisiana State Univ. 
34. 

Smithe, F. B. 1975. Naturalist’s Color Guide. Am. Mus. Nat. Hist., New York. 

Smithe, F. B. 1981. Naturalist’s Color Guide, part III. Am. Mus. Nat. Hist., New York. 


Address: Sebastian A. Lousada, Flag Hill Farm, P. O. Box 31, Vershire, VT 05079, 
U.S.A. Steve N. G. Howell, Point Reyes Bird Observatory, 4990 Shoreline 
Highway, Stinson Beach, CA 94970, U.S.A. 


© British Ornithologists’ Club 1997 


JF. W. Duckworth 210 Bull. B.O.C. 1997 117(3) 


Observations on a population of Jerdon’s 
Bushchat Saxicola jerdoni in the Mekong 
channel, Laos 


by FJ. W. Duckworth 
Received 4 September 1996 


Jerdon’s Bushchat Saxicola jerdoni ranges west from Vietnam to eastern 
India (King et al. 1975), but there are few recent records and little has 
been written specifically about it. It is regarded as globally 
near-threatened (Collar et al. 1994). There are several historical records 
from Laos, all from north of Vientiane (Delacour: & Jabouille 1927, 
Bangs & Van ‘Tyne 1931, Bourret 1943, David-Beaulieu 1944). 
Between 1950 and 1990, no new bird observations were reported from 
Laos, and during intensive fieldwork from 1992 to 1995 the species was 
not found at any of the numerous sites surveyed (Thewlis et al. 1996, in 
prep.), which were, however, mostly to the south of the historical 
records. In March 1996 a dense population was discovered breeding at 
the inflow of the Nam Sang river to the Mekong only 60 km upstream 
of Vientiane. Subsequent observations were made in June and July 
1996 at this site. The observations were made during an assessment of 
the conservation needs of the extensive area of cultivation and degraded 
forest to the north of Paksang (Duckworth 1996a, b) and relatively little 
time could be spent at the Mekong. 


Study area and methods 


The Nam Sang river enters the Mekong at Paksang (18°12'N, 102°09’E; 
170 m a.s.1.), where there are a number of sedimentary islands and bars 
(mainly sands and silts) amid the many extensive rocky outcrops from 
the river bed. Much of the sedimentary plain, which is exposed in the 
channel by the lower water levels during the dry season, supports a 
dense bushland, composed of few woody species, growing to a height of 
1—2 m. These bushes are submerged by turbulent water during the 
season of high flow, from mid-July to mid-October, after which water 
levels drop progressively; they are lowest during April. There are two 
main islands: Don Nou, at the mouth of the Nam Sang, and the much 
larger Don Sadok slightly downstream. Densely vegetated habitat 
extends 6 km along the river and in places exceeds 1 km in width; the 
total area estimated (from the Lao Service Geographique d’Etat 
1:100,000 maps) is 3-4 sq. km. The habitat on the downstream half of 
Don Sadok differs from the bushland on Don Nou, being dominated by 
extensive areas of rank grass and, in places, stands of mature trees. 
The area was visited in 1996 as follows: 14 February (12.00-—14.00 h); 
13 March (08.00-13.00h); 22 June (06.30-19.00h); and 15 July 
(06.30-18.30 h). Observations in February and March were limited to 
Don Nou. In June (when only half of the land exposed in March was 
still above water), work was concentrated on Don Nou but a brief boat 


F. W. Duckworth 211 Bull. B.O.C. 1997 117(3) 


trip was made around Don Sadok. In July the entire area of both 
islands remaining above water (about a tenth of that in March, and 
broken into many small islets) was surveyed. There was exceptionally 
little rain in early July 1996 and in many years there is probably 
negligible suitable habitat above water by mid-July. In both June and 
July, intensive observations were made on one morning on the dry land 
adjacent to the Mekong and for 1-2 km up the Nam Sang. This area 
is composed of rice paddies, dryland cultivation and regenerating 
scrub over abandoned cultivation amid fragmented and degraded 
Lagerstroemia-dominated mixed deciduous forest. 

Estimates of Jerdon’s Bushchat density were made in March by 
standing on four vantage points (all within 1 km of each other) and 
watching for 10-20 minutes to count the number of males in the 
viewable area, which itself was estimated as an are of x° (using a 
compass) with birds visible for ym (estimated visually; calibration 
elsewhere indicated that the observer could estimate distances across 
bushland of up to 500 feet [c. 150 m] within 20% accuracy). Visibility 
varied widely between vantage points and the method was crude 
because much of each area comprised bare rock, boulders or pools, 
none of which supported bushchats, and some birds were doubtless 
overlooked. 


Observations 


Counts of birds 

Males were much more visible than females (in total 38 adult males 
were seen but only 3 females) and so only males were used for 
population estimation; this difference in visibility between the sexes 
was also noted by Stanford & ‘Ticehurst (1938). On 13 March, 
combined density estimates in the four areas (of 2.5—15 ha) yielded 15 
territories in 25 ha. Bushland probably covered a third of these 25 ha. 
While walking between the vantage points, six further singing males 
were located incidentally. On 22 June, 13 males were found, mostly on 
Don Nou, but most suitable habitat on both islands remained 
unsurveyed. On 15 July a thorough survey of all islands found only 
four males, which probably represented at least half the males in the 
area. 

Most observations were made in the bushland, especially of Don 
Nou. There was not time to survey Don Sadok in March, but a male 
was seen from a boat in the extensive grass at the downstream end of 
this island in June, as well as several in the bushland of Don Sadok in 
June and July. No bushchats were seen in the extensive areas of trees, 
bare rock or sediment on the islands, up the Nam Sang river, or in the 
dryland habitats adjacent to the Mekong channel. 


Breeding cycle 

Most males observed in March were singing from sprays of 
vegetation projecting above the general level of the bushland, but they 
also perched for long periods in such positions without singing. They 
frequently dived to the base of an adjacent bush after a song bout. The 


¥. W. Duckworth 212 Bull. B.O.C. 1997 117(3) 


morning of observations was largely cloudy and song continued 
intermittently at high levels throughout the visit. The closest 
separation observed between males in March was 30m. In June and 
July, no singing was heard and, in June, two males were seen perched 
for many minutes only 6m apart without showing aggression; 
territorial behaviour seemed negligible, but the males still spent long 
periods (many minutes) perched prominently atop projecting twigs and 
grasses. A juvenile male with a full-grown tail was seen with a female 
on 22 June. 


Vocalisations 

The song, which, with that of Yellow-bellied Prinia Prinia 
flaviventris, was the predominant bird sound coming from the bushland 
in March, was a thin warble such as might be produced by a Sylvia 
warbler, but lacking any harsh or churring notes. The most frequent 
call was a plaintive, loud, single-note heeeeew; a single-note swearing 
rasp was occasionally given. 


Discussion 
Population and status in the island complex 

The territorial population on the islands in the Paksang area in 1996 
probably exceeded 100 pairs. The bushland occupied by the birds 
occurs in a fine mosaic with open sediment, rocks and water; this figure 
is calculated on the basis of three pairs per 5 ha of mosaic habitat (as 
above) of which there is at least 200 ha. If grassy areas of Don Sadok 
were also occupied, the total population might have reached 200 pairs. 

No bushchats were found in the special searches during June and 
July of the cultivation and scrub adjacent to the Mekong, nor were any 
seen in extensive surveys during February-March and June—July of 
such habitats over the area for 40 km to the north. The vegetation of 
some abandoned fields was of similar superficial physiognomy to the 
Mekong channel bushland, although the two habitats were composed of 
different plant species and the latter is a natural habitat while the 
former is a secondary regrowth in an area which was initially forested. 
A similar restriction, in that case of several species, to the vegetation of 
seasonally-inundated islands, with absence from adjacent dryland 
scrub, was also found in Amazonia by Rosenberg (1990) and Tye 
(1995). Both authors suggest that the distribution of the birds in 
question is restricted more by vegetation characteristics than by 
insularity per se; it is not possible to tell the relative importance of the 
two in determining Jerdon’s Bushchat distribution at Paksang as no 
natural bushland was found away from islands. 

During the months of highest flow, the Mekong inundates all habitat 
suitable for Jerdon’s Bushchat. The bushes are completely underwater, 
but it is this annual submersion that retains the vegetation as bushland. 
Most bushchats had left the channel by July, some probably by late 
June. It is not yet possible to suggest where the birds go when the 
Mekong is high, but the absence of observations in dry land vegetation 
at Paksang means that it is unlikely that they simply move from the 


J. W. Duckworth 213 Bull. B.O.C. 1997 117(3) 


channel into the adjacent vegetation, unless they change completely in 
behaviour and become highly skulking. In the context of observations 
elsewhere, and the low sighting rate of females on the islands, this latter 
is more possible that it might seem. Before extensive human 
modification to the environment, the bushchats would have gone to 
some natural habitat when the Mekong was high, and it is perhaps most 
likely that they still go there. It may be another form of natural 
bushland, as the allied Whinchat Saxicola rubetra, when making 
short-term stops on migration over lowland England, selects from the 
available hedgerows and scrub those which are most similar to its 
breeding habitat (Duckworth 1994). 


The lack of observations of the species on the February visit is of no 
significance as the two midday hours spent in the area were devoted to 
searching for waders and hirundines. 


Bird community of the Mekong channel at Paksang 

The distinct bird community in the Mekong channel at Paksang was 
typical of such a specialised island habitat with structurally simple 
vegetation: it had few species but a relatively high proportion were 
abundant, while some were rare or scarce elsewhere in the region. The 
bushland held high densities of Jerdon’s Bushchat, Plaintive Cuckoo 
Cacomantis merulinus, Greater Coucal Centropus sinensis, Streak-eared 
Bulbul Pycnonotus blanfordi, Oriental Magpie Robin Copsychus saularis, 
Common Tailorbird Orthotomus sutorius, Plain Prinia Prinia inornata 
and Yellow-bellied Prinia, with lower numbers of Blue-tailed Bee-eater 
Merops philippinus, Pied Bushchat Saxicola caprata, Bright-capped 
Cisticola Cisticola exilis and a few others. Like Jerdon’s Bushchat, the 
bulbul, bee-eater, Pied Bushchat, cisticola and both prinia species were 
extremely localised or unrecorded in the adjacent dry-land mosaic of 
cultivation, scrub and relict forest. Conversely, Grey-breasted Prinia 
Prinia hodgsoniu, Stripe-throated Bulbul Pycnonotus finlaysoni (and 
various other species) abounded up to the riverbank but were rare and 
unrecorded, respectively, in the channel. 


Some hillocks in the channel are always above water, except for flash 
floods, and their trees supported a depauperate derivative of the bird 
community of the adjacent land, including visits by Pompadour Green 
Pigeon Treron pompadora (at risk in Thailand and Laos; Treesucon & 
Round 1990, Thewlis et al. in prep.). Rocky outcrops within the 
channel hosted Wire-tailed Swallows Hirundo smithi1, and the earth 
cliffs Plain Martins Riparia paludicola; both probably bred and in July 
over 300 of the former and over 100 of the latter were observed. These 
are the largest numbers seen in Laos in recent years of both species, 
which are at risk in Thailand and have declined in Laos (Treesucon & 
Round 1990, Thewlis et al. in prep.). Sandbanks, open mud and pools 
held ducks, wagtails, Paddyfield Pipit Anthus rufulus and waders 
(migrant and resident), and notably small numbers of River Lapwing 
Vanellus duvaucelit which has declined in Laos (Duckworth et al. in 
press) and Small Pratincole Glareola lactea, which is at risk in Thailand 
(Treesucon & Round 1990). 


J. W. Duckworth 214 Bull. B.O.C. 1997 117(3) 


Comparison with other records of ferdon’s Bushchat 

These are the first records from Laos of Jerdon’s Bushchat for over 
50 years. The species is poorly known and previous information, from 
throughout its range, concerning habitat use has been conflicting. This 
section aims to summarise all published information about the species 
relevant to its population, distribution and habitat use. 

In Indochina, Jerdon’s Bushchat was considered to be a rare and 
localised resident on several high plateaux (Delacour & Jabouille 1931). 
In Laos, it was a common resident above 1400m in Tranninh, 
particularly at Ban Nonghet, a high valley (1500 m) dominated by rocks 
and tall herbs in place of the original forest and subject to frequent 
fires, frosts and fog (Delacour & Jabouille 1927, David-Beaulieu 1944). 
A specimen from Phou Khoum, a deforested mountain at 1600 m on 
the border of the provinces of Louangphabang and Tranninh, and 
lacking in ornithological interest (David-Beaulieu 1944), was received 
by Bourret (1943). A breeding female was taken on 12 May 1929 at Ban 
Muangyo in the far north of Laos at 2300 ft (750 m), an area of rice 
fields surrounded by good forest (Bangs & Van ‘Tyne 1931). 

The only other recent Lao record is of a male on 1 April 1996 in low 
bushes on a Mekong island 2 km downstream of the mouth of the Nam 
Ou (W. G. Robichaud verbally 1996). No Jerdon’s Bushchats were 
found during frequent observation in 1992-1996 on Don Chuan, a 
large island in the Mekong in Vientiane (‘Thewlis et al. 1996, in prep.), 
which however lacks the bushland of the Paksang region, nor were any 
seen on a 2-3 hour search, specifically for the species, of an extensive 
area of sand with patches of bushland around 'Thadua (25 km 
downstream of Vientiane) on 10 April 1996 (R. J. Tizard & R. J. 
Timmins verbally 1996); the species is so conspicuous that, if it is 
present, one may be certain of locating males during breeding season 
observations of river channel bushland (Stanford & Ticehurst 1938 also 
felt a similar certainty about seeing birds in this habitat). 

There are very few records from Vietnam: Kuroda (1917) and 
Delacour (1930) listed the species from Tonkin, at Lao Kay and Pakha 
respectively. Both are hilly areas, the latter being a quite deforested 
area at 4000 ft (1300 m). 

In Thailand, the species occurs only in the north, is very scarce and 
inhabits the tall grass of riverine floodplains; populations have been 
reduced by drainage and clearance, especially the burning of reedbeds 
for agriculture (Round 1988); Round (1983) specifies three sites. J. N. 
Dymond (in litt. 1996) observed the species at Chiang Saen in northern 
Thailand where small numbers of birds (maximum 3) were found in a 
narrow belt of tall riverside grass containing a few thorny bushes which 
the chats used as perches. All his observations were in January— 
February and the species was very skulking; no calls or song were 
heard. 

Only in Burma has occupation of river-island scrub been described 
previously: in the north bushchats were found with certainty where 
wild roses (called Rosa bracteata in Smythies 1986) grew on the sandy 
islands of the larger river valleys, including at Sinho (where it was 
probably breeding in May) and on the western edge of the Indawygi 


F. W. Duckworth 215 Bull. B.O.C. 1997 117(3) 


lake (Stanford & Ticehurst 1938). In Bhamo it bred in long grass and 
briars in the Irrawaddy valley in April-May (Harington 1905, 1909). 
Otherwise, its status in the country caused some controversy: Stanford 
& Ticehurst (1938) felt that the species occurred only in vast grass 
plains at low altitudes and disagreed with a former statement that it was 
a hill bird. Blanford (1870) found it in elephant-grass in the Irrawaddy 
valley, and Wickham (1929) found it only in this valley, in the northern 
part of the country. Oates (1883) considered that although probably 
resident, it was extremely rare in Burma, quoting only Blanford’s 
(1870) record and some from Bhamo. Stanford & Ticehurst (1935) shot 
only one in lower Burma in over three years, in kaing grass in 
Leikchuang, and agreed with Oates’s assessment of it as extremely rare 
in that part of Burma. Up to nine were recorded on three 
January—February visits in 1973-1982 to the reeds around Inle Lake 
(King 1983). Birds are still common (at least 10 males were seen in one 
day) at this lake, in 3-4 m high reeds at the lake’s edge and in the huge 
floating mass of reeds on the lake itself; birds were observed feeding 
in areas of aquatic crops and floating vegetable gardens amid this mass 
(F. R. Lambert in litt. 1996). 

In India it was: common all over the Brahmaputra basin in thick 
vegetation (particularly grass hedges) along streams or wet ditches, and 
one of the commonest birds in Manipur, including the suburbs of the 
capital Imphal, but it was never seen in the hills (Hume 1888); very 
common in North Cachar in cold weather, with a few remaining to 
breed on the higher hills (Baker 1894-1901); a common resident in the 
Khasia hills, and common on the adjacent plains in cold weather, but 
ascending above 3000 ft (900 m) in the breeding season (presumably 
about March) (Baker 1907); resident in parts of Upper Assam, 
occurring throughout the plains in the cold season (October to 
February) and confined to reed and grass adjacent to rivers (Stevens 
1914-1915); an abundant breeder (laying eggs in April—May) in vast 
plains of sun grass on the north bank of the Brahmaputra in Lakhimpur 
district, and occurring at the base of the foothills in this district, in 
smaller numbers west to the grass plains of North Kamrup district, and 
also in the upland grass plains of the Chin hills (Baker 1924). Baker’s 
(1933) summary of its status in India (which was not added to by Ali & 
Ripley 1973) recorded it from vast expanses of various tall grasses in 
the plains at the base of the Himalayas, extending into upland grass 
plains and foothills up to 2500 ft (700 m). It was particularly common 
in Assam in many places in Sibsagar and North Lakhimpur, where it 
bred in February—April in grasslands and grassy sandbanks in rivers 
that were flooded towards the end of that month by Himalayan 
meltwater. 

There are very few recent records for the Indian subcontinent, where 
most areas of long grass in the wet lowlands, from which the majority 
of records listed above come, have been modified for cultivation 
(T. P. Inskipp verbally 1996). Choudhury (in press) recorded singles 
(over several years) each in November, March and May, in grass and 
reed along river or channel banks in the Dibru-Saikhawa Wildlife 
Sanctuary, Assam. This area is on the Brahmaputra plains just 


JF. W. Duckworth 216 Bull. B.O.C. 1997 117(3) 


north-east of the area where Stevens (1914-1915) had previously 
recorded it regularly. ‘There are only few records from Kaziranga 
National Park, Assam, an area of extensive grassland which seems, 
from previous descriptions of habitat occupied by the species in 
Assam and Burma, to be ideal for the species (T. P. Inskipp verbally 
1996). 

There are only three recent published records from Bangladesh, all 
from tea estates or grassland in the northeast in December—February 
(Thompson et al. 1993); the only previous records from the country 
cited in this source (from Godwin-Austen 1870) give no information on 
status or habitat use. 

The only records from Nepal appear to be from the Kosi barrage in 
the east in May—June 1975-1976 (Inskipp & Inskipp 1991) and a few in 
the early 1990s from the west of the country (N. J. Collar verbally 
1996); all were in extensive grassland. 

‘There are few records from Yunnan: a male along the Namting river 
in February 1917 (Rothschild 1926), four specimens collected in Luxi 
and Gengma counties in the 1960s and a few sightings in Menglun, 
Xishuangbanna, in April 1994 during a two month survey (Han 
Lianxian in litt. 1996). The latter birds were all within 100 m of rivers. 

In summary, although the records from the Mekong channel at 
Paksang are in a different habitat from the deforested hills in which it 
was previously recorded in Laos, records outside Indochina come 
mainly from lowland grass and scrub, usually close to water. The 
most frequently noted vegetation, extensive tall floodplain grassland, 
is rare or perhaps absent from the Mekong in north-central Laos. It 
might be that the bushchats at Paksang are inhabiting a sub-optimal 
area, following loss of a more favoured grassland (J. C. Eames zn Mitt. 
1996); however, if this were true it is unlikely that the density on the 
islands would be so high, or that the bird would be so localised to 
one habitat. Furthermore, occurrence in both lowland river valley 
scrub and higher-altitude hill scrub has previously been documented 
only in Burma. In some areas of India and Burma, the bird was 
found at low altitudes only in winter, with several authors noting a 
move to higher altitudes to breed, although the most comprehensive 
summary referred to breeding in seasonally flooded areas, as 1s 
Paksang. In general the bird’s occurrence seemed to be patchy, with 
areas of local abundance amid extensive areas where the bird was 
scarce. 


Threats to, and conservation needs of, the species in Laos 

In contrast to forest, direct human exploitation of the island 
vegetation does not currently pose a threat to the habitat. It currently 
supports low levels of stock grazing and a major increase in this might 
affect the vegetation structure and in turn the bushchats. Salad and 
melons are grown during the dry season on the areas of extensive open 
sediment not used by the bushchats. Currently only a small proportion 
of the area suitable for gardens is used as such and no bushland is 
cleared for this purpose. The shrub Mimosa pudica, which thrives in 
disturbed areas, has taken over huge areas of lowland northern 


¥. W. Duckworth 217 Bull. B.O.C. 1997 117(3) 
Thailand (P. D. Round zn litt. 1996). Although it has not colonised the 


Paksang islands, if their natural vegetation were to be cleared or 
overgrazed for any of the above reasons, regrowth might involve this 
invasive species, whose suitability for Jerdon’s Bushchat is not known. 

The heavy use of the area for fishing results in high levels of 
disturbance which are detrimental for River Lapwings (Duckworth 
et al. in press) but are unlikely to affect the bushchats, which seemed 
unconcerned by observers even within 10 m. Most small boys in Laos 
habitually kill birds with catapults and the habit of Jerdon’s Bushchats 
of perching prominently on low vegetation makes them an easy 
potential target; but as the bushland can be reached only by boat, the 
threat from boys is negligible. 

Potential threats which are more serious come from eleinges in the 
sediment flow dynamics of the Mekong as the bushland grows upon 
river-borne sediment. If substantial quantities of sediment are removed 
from the river, or trapped by barriers, water with an unnaturally low 
sediment load will wash away, without replenishment, downstream 
sand and silt deposits (Brooker 1981). If this were to happen upstream 
of Paksang, it would result in the reduction or even loss of vegetation. 
Intensive extraction of sediment during the dry season from the 
Mekong in Vientiane since 1994 has caused a major shrinkage of Don 
Chuan, a large island formerly supporting extensive vegetation. During 
1996, similar heavy extraction was observed upstream of Vientiane 
halfway to Paksang. It is not clear if the sustainability of this practice, 
which was initiated only in the last few years, has been considered by 
those engaged in it, and it may be that as deposits close to Vientiane 
become depleted, extraction sites will move upstream. The suitability 
of the Paksang region for bushchats (the sediment of much of which, 
especially the areas supporting bushland, is too interspersed with 
rocky outcrops to make extraction financially viable from the site 
itself) would be reduced if substantial extraction were to occur 
upstream of it. 

A further potential threat is the proposed chain of cross-Mekong 
hydropower dams which, if built, would disrupt seriously the ecology 
of the entire basin (Roberts 1995), probably including the sediment 
flow, as each dam traps sediment in its headpond. 

The urgency for conservation action for this species is thus lower 
than for forest birds (see Thewlis et al. in prep.) and its precise level is 
difficult to assess because of the paucity of information on the bird in 
Laos, as elsewhere in its range. Further survey is thus the highest 
priority action. Visits to Paksang are needed to clarify the extent of 
occupied habitat, the population and the seasonal status of Jerdon’s 
Bushchats there between August and February. The extent of occupied 
habitat along the rest of the Mekong and its major tributaries in Laos 
should be investigated. The areas and habitats used by the birds from 
Paksang during the peak of the wet season need to be established; they 
may have been modified or reduced by human activity. The montane 
areas of secondary grassland used by the birds in Tranninh (now Xieng 
Khouang province) need to be revisited and the bird’s current status 
there assessed. 


¥. W. Duckworth 218 Bull. B.O.C. 1997 117(3) 


Summary 


A dense population of the globally near-threatened and little-known Jerdon’s Bushchat 
Saxicola jerdoni (totalling 100-200 pairs over 6sq. km) was discovered in March 1996 
breeding amongst a complex of scrub dominated islands in the Mekong channel at 
Paksang, 60 km upstream of Vientiane. Small numbers of birds were still present in mid 
July when most of the breeding area had been submerged by the seasonally rising river 
water. The remainder of the birds could not be located on the adjacent dry land and 
appear to disperse out of the area. This habitat differs greatly from that recorded in Laos 
during the only previous observations, made during 1925-1945, but such breeding 
habitat has been described previously in Burma. The species is probably under no 
imminent threat at the site unless the rapidly expanding practice of removing sediment 
from the channel during the dry season extends upstream of the site. Further surveys are 
urged, both in the Mekong and its major tributaries, and in the montane scrub in the 
north of Laos which was previously reported to hold the species. These are particularly 
important in view of the paucity of recent records from elsewhere in the species’ range. 


Acknowledgements 


Permission to conduct the survey was granted by the District Governor of Sangthong, 
Mr Kenta Kongbounma. Advice and assistance was received from the director of the 
Vientiane Forestry College, Mr Soukkongseng Saignaleuth, the Lao-German Forestry 
Teamleader, Dr H.-M. Brechtel, Walter Ehrhardt, Tim Inskipp, Bill Robichaud, Craig 
Robson, Rob Timmins, Rob Tizard and Hilary Tye. Nigel Collar, Mike Crosby, Nick 
Dymond, Jonathan Eames, Han Lianxian (Kunming Institute of Zoology), Frank 
Lambert, Philip Round and Richard Thewlis kindly supplied details of other records or 
commented on the manuscript. Mr Chanthaphone Phon-asa worked extremely hard and 
capably during the survey. The survey on which these observations were made was 
conducted under contract to the Lao-German Forestry Team project for the Promotion 
of Education at Vientiane Forestry College with funding from GTZ. 


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Baker, E. C. S. 1894-1901. Birds of North Cachar, part 3. 7. Bombay Nat. Hist. Soc. 9: 
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Baker, E. C. S. 1907. Birds of the Khasia hills. 7. Bombay Nat. Hist. Soc. 17: 783-795; 
957-975. 

Baker, E. C. S. 1924. The Fauna of British India including Ceylon and Burma. Birds vol. 2. 
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Baker, E. C. S. 1933. The Nidification of Birds of the Indian Empire, vol. 2. Taylor and 
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Bangs, O. & Van Tyne, J. 1931. Birds of the Kelley-Roosevelt’s expedition to French 
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Blandford, W. T. 1870. A list of birds obtained in the Irawadi valley around Ava, Thayet, 
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Bourret, R. 1943. Liste des oiseaux récemment entrés dans les collections du Laboratoire 
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Brooker, M. P. 1981. The impact of impoundments on the downstream fisheries and 
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Choudhury, A. in press. Bird survey in Dibru-Saikawa Wildlife Sanctuary, Assam, 
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Collar, N. J., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to Watch 2: the world list of 
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UK 

David-Beaulieu, A. 1944. Les oiseaux du Tranninh. Université Indochinoise, Hanoi. 

Delacour, J. 1930. On the birds collected during the fifth expedition to French 
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Delacour, J. & Jabouille, P. 1927. Recherches ornithologiques dans le Provinces du Tranninh 
(Laos), de Thua-Thien et de Kontoum (Annam) et quelques autres régions de 


JF. W. Duckworth 219 Bull. B.O.C. 1997 117(3) 


l’Indochine Francaise. Archives d?Histoire Naturelle. Société Nationale 
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Delacour, J. & Jabouille, P. 1931. Les otseaux de Il’Indochine Frangaise Vols I-IV. 
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Duckworth, J. W. 1994. Habitat selection by migrant Redstarts Phoenicurus phoenicurus 
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Duckworth, J. W. 1996a. Conservation concerns in the Vientiane Forestry College Training 
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Duckworth, J. W. 1996b. Bird and mammal records from the Sangthong District, 
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A. Aleixo 220 Bull. B.O.C. 1997 117(3) 


Tye, H. 1995. Ecology of the bird community of regenerating forest habitats in lowland 
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Address: J. W. Duckworth, East Redham Farm, Pilning, Bristol BS12 3JG, U.K. 
© British Ornithologists’ Club 1997 


Range extension of the Large-headed Flatbill 
Ramphotrigon megacephala with comments on 
its distribution 


by Alexandre Aleixo 
Received 13 March 1996 


The Large-headed Flatbill Ramphotrigon megacephala is patchily 
distributed in the forests of the Andes (western and central Venezuela, 
eastern and northern Colombia on east and west slopes, eastern 
Ecuador), western Amazonia (southeastern Peru, northern Bolivia, 
western Amazonian Brazil), humid montane Atlantic forest in 
southeastern Brazil (eastern Minas Gerais, Rio de Janeiro south to Sao 
Paulo), subtropical deciduous forests of eastern Paraguay (Alto Parana, 
Canendiyu), and northeastern Argentina in Misiones (Ridgely & Tudor 
1994, Hayes 1995, Pacheco 1995). This inconspicuous small flycatcher 
(13 cm; 13-17 g) seems to be largely linked to bamboo patches in all the 
regions where it occurs, perching low in the dense interior of the 
thickets, from which it sallies out for insects on bamboo foliage and 
stems (Parker 1982, 1984, pers. obs). Though difficult to observe (it 
remains motionless for long periods in the dense understory), it is 
readily detected when calling a ventriloquial disyllabic whu-hoo, mainly 
in early morning and late afternoon (Parker 1984, pers. obs). 

On 22 October 1995, at 06.55 hrs, I tape-recorded an individual of 
Ramphotrigon megacephala (ALPA 21/10, deposited in the Library of 
Neotropical Sounds of the State University of Campinas—U NICAMP 
[ASN]) in a riparian forest with an understory of dense bamboo 
thickets, about 120km northwest of the town of Barra do Gargas 
(15°53’S, 52°15'W, on the western bank of the upper Araguaia river in 
southeastern Mato Grosso State, central Brazil). The riparian forest 
was on the margin of the Mortes river at an altitude of about 350 m in 
the Xavante Indian reserve of SAo Marcos. The dominant vegetation of 
the area is open cerrado (savanna-like vegetation), with riparian forests 
having a variable amount of understory bamboo cover. 

The nearest known locality where R. megacephala has been reported 
is Alta Floresta in northern (Amazonian) Mato Grosso State (Ridgely & 
Tudor 1994), about 700km northwest of the Sao Marcos Indian 
reserve. I compared the voice of the bird from Sao Marcos reserve with 
two other birds recorded in Ji-Parana (Rond6énia State, Amazonian 


A. Aleixo 221 Ball. BOC. 199F 1178) 


FREQUENCY . 
(kHz) 
py 


2 
OK 
EEE 
0.5s$ 
B 
8 


FREQUENCY 
(kHz) 
» 


Ls) 


FREQUENCY 
(kHz) 


oe 
0.5s 


TIME (S) 


Figure 1. Sonagrams of the song of three populations of the Large-headed Flatbill 
Ramphotrigon megacephala in Brazil. A, Ji-Parana, Rondénia State, Amazonia (JV 
495/5b); B, Barra do Garcas, Mato Grosso State, central Brazil (ALPA 21/10); C, Parque 
Estadual Intervales, Sao Paulo State, southeastern Brazil (ALPA 26/1+2). All recordings 
are deposited in the Libary of Neotropical Sounds (ASN) at the Bioacoustical Laboratory 
of the Universidade Estadual de Campinas-UNICAMP. Sonagrams were made on a 
MacIntosh Classic coupled: to a MacRecorder Sound System 2.0.5. 


Brazil; JV 495/5b-ASN) and Parque Estadual Intervales, Capao Bonito 
(southern Sao Paulo State, Brazil; ALPA 26/1+2-ASN). The song 
structure of the Sao Marcos bird (the disyllabic whu-hoo) is very similar 
to that of the birds from Amazonia (Rond6nia) and southeastern Brazil 
(Fig. 1). Though a larger sample size is needed, preliminary measures 
of the duration of the notes of the song of the SAo Marcos bird indicate 
its closer relationship to Amazonian birds (Table 1). 

The range extension here reported fills a gap in the spotty 
distribution pattern of R. megacephala, which until 1939 was known 
only from a few specimens from southeastern Brazil and adjacent 


A. Aleixo 222 Bull. B.O.C. 1997 117(3) 


TABLE 1 
Duration of the notes and interval between the two notes (in seconds) of the song of three 
populations of the Large-headed Flatbill Ramphotrigon megacephala in Brazil: Amazonia 
(JV 495/5b, Ji-Parana, Rondénia State), Central Brazil (ALPA 21/10, Barra do Gargas, 
Mato Grosso State) and Southeastern Brazil (ALPA 26/1+2, Capao Bonito, Sa0 Paulo 
State). Sounds were analysed on a MacIntosh Classic coupled to a MacRecorder Sound 


System 2.0.5. 
First note Second note Interval 
Amazonia 0.309 0.166 Q723 
Central Brazil 0.283 0.165 0.26 
Southeastern Brazil 0.211 0.133 0.28 


Argentina and Paraguay (Zimmer 1939). Further research has revealed 
a broader though very local distribution (Novaes 1960, Parker 1984, 
Ridgely & Tudor 1994, Pacheco 1995). The population here reported 
of R. megacephala in central Brazil, a region of dry open vegetation 
which separates two large humid forest tracts (Atlantic forests and 
Amazonia) and two distinct populations of the Large-headed Flatbill 
(R. m. megacephala and R. m. boliviana, respectively), may be inter- 
preted as relictual. The new locality for R. megacephala lies approxi- 
mately at equal distance between southwestern Amazonia and 
southeastern Brazil, two regions known to harbour a very distinct and 
specialist avifauna associated with bamboo thickets (Parker 1982, 
Pierpont & Fitzpatrick 1983, Rodrigues et al. 1994). This new record 
suggests that R. megacephala may have had a wider distribution in the 
past, being now confined to regions where its habitat still remains. 
Relictual bamboo understory forests in central Brazil have not 
previously been reported in the literature (Sick 1955, Goodland 1971), 
and may be very locally distributed. Prance & Brown (1987) 
hypothesized that during the Pleistocene the transitional forest (which 
includes the bamboo forest type) was more extensive but was replaced 
more recently by rain forest in Amazonian lowlands and by cerrado on 
the central Brazilian Plateau. After the reduction of the transitional 
forest due mainly to climatic changes, species peculiar to it would have 
a relatively restricted and fragmented distribution (Prance & Brown 
1987), a hypothesis for which this record of the Large-headed Flatbill 
provides support. Further ornithological investigation in these relictual 
riparian forests with a dense understory of bamboo will probably lead 
to other range extensions of forest birds closely associated with 
bamboos and previously unknown on the central Brazilian shield. 


Acknowledgements 


FAPESP (Fundacao de Amparo a Pesquisa do Estado de Sao Paulo) provided financial 
support for the avifaunal survey of the Sao Marcos Indian reserve as part of the study on 
bird classification by Xavante Indians, conducted by the anthropologist Eduardo Carrara, 
who helped in many ways. I am deeply indebted to Jacques Vielliard, Keith S. Brown Jr. 
and Marco Aurélio Pizo for helpful comments and review of the manuscript. 
Célio F. D. Haddad provided invaluable assistance with sound analysis and sonagrams at 


S. K. Herzog et al. 223 Bull. B.O.C. 1997 117(3) 


the Bioacoustical Laboratory of UNESP-Rio Claro (sponsored by FAPESP, grant no. 
94/5634-5). The Conselho Nacional de Desenvolvimento Cientifico e Tecnoldgico 
(CNPq) and FMB financed materials and equipment at the Bioacoustical Laboratory of 
UNICAMP. I also thank FAPESP for a research fellowship and the Xavante Indian 
community of Namunkura for their hospitality. 


References: 

Goodland, R. 1971. A physiognomic analysis of the cerrado vegetation of central Brazil. 
Jj. Ecol. 59: 411-419. 

Hayes, F. H. 1995. Status, Distribution and Biogeography of the Birds of Paraguay. 
Monographs in Field Ornithology no. 1. American Birding Association. 

Novaes, F. C. 1960. Sobre Ramphotrigon megacephala (Swainson) (‘Tyrannidae, Aves). 
Rev. Bras. Biol. 20: 217-221. 

Pacheco, J. F. 1995. New distributional records for some birds from varzea forest at 
Mamirana Reserve, Western Brazilian Amazonia. Ararajuba 3: 83-87. 

Parker, T. A., III. 1982. Observations of some unusual rainforest and marsh birds in 
southeastern Peru. Wilson Bull. 94: 477-493. 

Parker, T. A., III. 1984. Notes on the behavior of Ramphotrigon flycatchers. Auk 101: 
186-188. 

Pierpont, N. & Fitzpatrick, J. 1983. Specific status and behavior of Cymbilaimus 
sanctaemariae, the bamboo antshrike, from southwestern Amazonia. Auk 100: 
645-652. 

Prance, G. T. & Brown, K. S., Jr. 1987. Soils and vegetation. Pp. 28-45 in T. C. 
Whitmore & G. T. Prance (eds), Biogeography and Quaternary History in Tropical 
America. Clarendon Press, Oxford. 

Ridgely, R. S. & Tudor, G. 1994. The Birds of South America. Vol. 2. The Suboscine 
Passerines. Oxford Univ. Press. 

Rodrigues, M., Alvares, S. M. R. & Machado, C. G. 1994. Foraging behavior of the 
White-collared Foliage-gleaner (Anabazenops fuscus), a bamboo specialist. Ornitolo- 
gia Neotropical 5: 65-67. 

Sick, H. 1955. O aspecto fitofisiondmico da paisagem do médio Rio das Mortes, Mato 
Grosso, e a avifauna da regiao. Arg. Mus. Nac. Rio de F. 42: 541-576. 

Zimmer, J. T. 1939. Studies of Peruvian birds no. 33. The genera Tolmomyias and 
Rhynchocyclus with further notes on Ramphotrigon. Amer. Mus. Novit. 1045. 


Address: Alexandre Aleixo, Departamento de Zoologia, Universidade Estadual de 
Campinas, Caixa Postal 6109, Cep 13083-970, Campinas-SP, Brazil. 


© British Ornithologists’ Club 1997 


Distributional notes on birds of Andean dry 
forests in Bolivia 


by Sebastian K. Herzog, Michael Kessler, Sjoerd Mayer & 
Stefan Hohnwald 


Received 15 May 1996 


From 20 May to 19 October 1995, an ornithological and botanical 
expedition conducted by the Foundation for Tropical Research and 
Exploration (TREX), in collaboration with the Herbario Nacional de 
Bolivia (HNB) and the Coleccién Boliviana de Fauna (CBF), visited 12 
Andean dry forest localities (i.e. forest mainly composed of 
drought-deciduous trees) throughout Bolivia. The avifauna of one of 
these sites (Inquisivi, site 6) had previously been surveyed in 1993, 
1994 and 1995 by SM, who did not participate in the TREX 


S. K. Herzog et al. 294 0 Bull. B.O.C. 1997 117(3) 


expedition, and his data are included in this paper. We present details 
on noteworthy ornithological results, including some notes on a species 
new to science (Cranioleuca, unnamed species), the first observations 
of Lemon-browed Flycatcher Conopias cinchoneti in Bolivia, 29 new 
departmental records, 12 new altitudinal records and notes on two 
threatened bird species. Remsen & Traylor (1989) and A birdlist of 
Bolivia (Armonia 1995) compiled and published by Asociacién 
Armonia, the Bolivian representative of BirdLife International, were 
used as references for all new departmental records listed below. New 
altitudinal records are mainly based on Fyeldsa & Krabbe (1990), 
Ridgely & Tudor (1989, 1994) and Armonia (1995). For all survey sites 
except Rio Pilcomayo (site 12), which was visited for only 2 days, lists 
of all species observed are presented (see Appendix). Tape-recordings 
of individual birds and dawn choruses were made at sites 2—6, 8 and 11 
and will be housed in the Library of Natural Sounds, Laboratory of 
Ornithology, Cornell University. 

Throughout Bolivia, dry forests at medium to high altitude (1500- 
3500 m) have been severely degraded or locally completely destroyed, 
mainly due to overgrazing, the excessive use of fire and the extraction of 
timber (especially for the production of charcoal). ‘Thus, in many areas 
(e.g. Torotoro, upper Consata Valley, upper tributaries of Rio Grande 
and Rio Pilcomayo) most dry forest is characterized by a low, open canopy 
and shows little to no tree regeneration. The final stages of degradation are 
Dodonaea-covered slopes or barren badlands. Consequently, conservation 
measures are urgently needed, especially since these forests hold a number 
of bird species that have small ranges (J. V. Remsen 2m Uitt.). 

In contrast, foothill dry forest dominated by Anadenanthera is still 
widespread, and, while little of this habitat can be regarded as pristine 
(in fact, pre-Incan terraces are commonly found), the ecosystem as such 
seems to be comparatively healthy. Fortunately, the most important 
area of foothill dry forest was recently included in Parque Nacional 
Alto Madidi, but other significant areas currently lacking any protective 
status are found along the lower Rio Grande in the Masicuri region and 
southward along the Cordillera de los Milagros, including our study 
area at Rio Azero. 


Survey sites: 

(1) Consata, depto. La Paz, provs. Saavedra and Munecas, from 
c. 3km (15°24’S, 68°31’W) to 18 km (15°26’S, 68°34°W) SW Consata 
along the road to Sorata; 30 May-3 June; 6-15m tall, 30-85% 
deciduous dry forest dominated by Anadenanthera colubrina (1000- 
1400 m); further downriver the forest graded into disturbed evergreen 
forest, further upriver into dry scrub. 

(2) Yolosillas, depto. La Paz, prov. Nor Yungas, c. 4km N Yolosa 
on the road to Caranavi (16°12’S, 67°45’W); 20-23 May, 8 Oct; small 
(1-5 ha), isolated patches of degraded (logging) dry forest (8-12 m tall, 
c. 50% deciduous, dominated by Anadenanthera colubrina) on steep 
slopes near the Rio Coroico (1000m) and on N-facing slopes 
(1100-1300 m); slopes facing W, E and S and most areas adjacent to the 
river supported degraded (logging, burning) evergreen forest. 


S. K. Herzog et al. 225 Bull. B.O.C. 1997 117(3) 


(3) Las Mercedes, depto. La Paz, prov. Sud Yungas, from 6 km 
(16°17'S, 67°23'W, 1300 m) to 27 km (16°13’S, 67°13’W, 800 m) E Villa 
Barrientos along road to La Asunta; 2—7 Oct; slightly degraded forest 
(10-15 m tall, c. 80% deciduous, dominated by Anadenanthera sp., 
Cereus sp., several Leguminosae) from Rio Boopi (800-850 m) and Rio 
Tamampaya (850-1000 m) up to c. 1200 m, above which dry forest was 
originally replaced by more humid forest but now mostly cleared for 
agriculture; c. 32 km E Villa Barrientos dry forest started to grade into 
humid foothill forest. 

(4) Miguillas, depto. La Paz, prov. Inquisivi, 4.5km NNW 
Miguillas (16°33’S, 67°22'W); 21-25 Sept; moderately. to, in part, 
severely degraded (logging, burning, grazing) dry forest (8-10 m tall, 
c. 70% deciduous, with Anadenanthera sp., Acacia macracantha, Cereus 
sp.) from Rio La Paz (1100 m) up to 1600-1700 m, above which the 
forest was largely replaced by pastures; shghtly more humid forest 
(10-12'm tall, c. 50% deciduous) in a shallow, c. 50 m wide, S-facing 
ravine. 

(5) Huara, depto. La Paz, prov. Sud Yungas, from the confluence of 
Rios La Paz and Jucumarini (16°37’'S, 67°28’W, 1250 m) to c. 3 km 
ENE Rancho Cieneguillas (16°34’S, 67°25’W, 1500 m); 27 Sept—2 Oct; 
moderately degraded dry forest (6—12 m tall, on steep, eroded slopes 
replaced by open scrub, 80—90% deciduous, dominated by several 
Leguminosae, Schinopsis sp., with Tillandsia durati abundant) from 
Rio La Paz (1200-1250 m) up to 1800-1900 m, above which the 
forest had largely been cleared; evergreen forest along the Rio 
Jucumarini (c. 12 m tall, 5-15 m on each side of the river, dominated 
by Cecropia sp., Solanum sp., Anadenanthera sp., Inga sp., Tessaria 
integrifolia, Piper sp.) interspersed with gravel bars and open scrub 
vegetation. 

(6) Inquisivi, depto. La Paz, prov. Inquisivi, slope N Inquisivi 
(16°54’S, 67°09°W) from 2500m down to Rio Khatu at 2050m 
(c. 2.5km N Inquisivi); 14-21 Sept, 17-18 Oct (TREX expedition); 
13-16 and 23-28 Dec 1993, 27 Jan—2 Feb 1994, 3-6 Jan 1995 (SM); 
moderately degraded (grazing, logging) dry forest (8-10 m tall, c. 95% 
deciduous, with Schinus molle, Schinopsis haenkeana, Ceiba sp.), largely 
replaced by agricultural fields and plantations of Eucalyptus globulus 
above 2500 m; a 50-80 m wide band of c. 20% deciduous forest along a 
ravine from c. 2500 m down to Rio Khatu; a small, severely degraded 
(logging, grazing) patch of evergreen forest and scrub at Rio Khatu 
(10 m wide stretch of c. 12 m tall forest along the river dominated by 
Erythrina sp., adjacent scrub up to 5 m tall and dominated by Piper sp., 
Acacia macracantha and several Compositae). 

(7) Rio Caine, depto. Potosi, prov. Charcas, and _ depto. 
Cochabamba, prov. Arce (18°06’'S, 65°46’'W to 17°53’S, 65°55’W); 
11-14 June; extensive areas of severely degraded (logging, grazing, 
burning) dry forest (c. 10m tall, 85% deciduous, originally domi- 
nated by Schinopsis haenkeana and Aspidosperma quebracho-blanco, 
now mostly by Prosopis kuntzei and Acacia spp.) on slopes at 2100- 
2400 m; extensive agricultural areas in the flat river valley (2050- 
2100 m) with very few trees (mostly P. kuntzei, A. quebracho-blanco, 


S.K. Herzog et al. 226 Bull. B.O.C. 1997 117(3) 


Capparis sp.) as well as Opuntia sulphurea and Puya sp. in overgrazed 
areas. 

(8) San Juan del Potrero, depto. Santa Cruz, prov. Florida, 5.5 km 
SE San Juan del Potrero (17°59’S, 64°15’W); 3-8 Sept; moderately 
degraded (grazing) dry forest (5—8 m tall, c. 80% deciduous, dominated 
by Aspidosperma quebracho-blanco, several Leguminosae and Cereus 
spp.) covering most of the area from seasonally dry creeks (where the 
forest was slightly taller and less deciduous) at 1500 m up to hill tops at 
1950 m. 

(9) Novillero, depto. Cochabamba, prov. Campero, from c. 3 km 
(18°18'S, 65°15°W, 2400 m) to 22 km (18°13’S, 65°18’W, 2800 m) NW 
Novillero along the road to Santiago; 16—20 June; degraded (logging, 
grazing) dry forest (c. 8 m tall, 80% deciduous, dominated by Tipuana 
tipu) grading into more humid forest (to 18m tall, 25% deciduous, 
including many Myrtaceae) on S-facing slopes at higher elevations; 
large areas cleared and dominated by scrub of Baccharis spp., 
Maiunthostachys sp., Tecoma sp., etc.; valley bottoms mostly converted to 
agriculture, only scattered patches of degraded evergreen forest 
(10-15 m tall, c. 15% deciduous) in ravines. 

(10) Masicuri, depto. Santa Cruz, prov. Vallegrande, from. the 
confluence of Rio Masicuri and Rio Grande (19°04’S, 63°41°W, 500 m) 
to the vicinity of Masicuri (18°49'S, 63°48'W, 800m); 9-14 July; 
slightly to severely degraded (logging, grazing) dry forest (8-15 m 
tall, 60-85% deciduous, dominated by several Leguminosae incl. 
Anadenanthera colubrina), gradually becoming more humid with 
increasing elevation until replaced by slightly to moderately degraded 
evergreen forest (c. 20 m tall, 10—20% deciduous) at about 800 m; most 
of the 0.5-1.5km wide valley bottom converted to pastures and 
plantations with few remaining patches of degraded (logging, grazing) 
evergreen gallery forest (to 30 m tall, c. 30% deciduous, originally less 
deciduous). 

(11) Rio Azero, depto. Chuquisaca, prov. Siles, from ¢. 15 km NW 
Rio Azero (19°32’S, 64°10°W, 1500 m) along the road to Padilla to 
35km SW Rio Azero (19°47'S, 64°02'W, 1200 m) along the road 
to Monteagudo (altitude at Rio Azero 1000 m); 27 June—4 July; 
large tracts of slightly disturbed (some grazing and logging) dry 
forest (15-20m tall, 40-80% deciduous, diverse with mumerous 
Leguminosae); slightly to severely degraded (logging) evergreen forest 
(to 25 m tall, c. 10% deciduous) along the Rio Azero and in humid 
ravines. 

(12) Rio Pilcomayo, depto. Potosi, prov. Linares, and depto. 
Chuquisaca, prov. Yamparaez, from Oron Kkota (19°34’'S, 64°51'W, 
2050 m) to 5km SW Icla (19°23’S, 64°48’W, 2450 m); 22-24 June; 
arid area with scattered patches of severely degraded dry forest 
(c. 7m tall, 95% deciduous, dominated by Schinopsis haenkeana, 
Tipuana tipu and Aspidosperma quebracho-blanco) at higher elevations; 
desert scrub (with Neoraimondia herzogiana) and degraded riverine 
forest (5-12 m tall, c. 50% deciduous, dominated by Aspidosperma 
quebracho-blanco, Fatropha hieronymi and various columnar cacti) at 
lower elevations. 


S. K. Herzog et al. 227 Bull. B.O.C. 1997 117(3) 


Species accounts 
BAND-TAILED PIGEON Columba fasciata 


This species was observed at 1500 m at Huara by SKH and MK, and 
at 800 m along the Rio Boopi at Las Mercedes by SKH, well below its 
usual elevational range (Fjeldsa & Krabbe 1990, Armonia 1995). 


MAROON-CHESTED GROUND-DOVE Claravis mondetoura 

Up to 5 individuals (3 males, 2 females) of this poorly known species 
were observed by SKH and SH on 3, 4 and 6 Oct within 2.5 km of road 
along the Rio Boopi and Las Mercedes, 500m below its known 
elevational range (Fjeldsa & Krabbe 1990, Armonia 1995). Surpris- 
ingly, all birds were observed in dry forest. Any humid forest with 
bamboo thickets, the habitat the species usually is associated with 
(Hilty & Brown 1986), was located on the slopes at least 400 m above 
the dry forest. 


RED-FRONTED MACAW Ara rubrogenys 

To increase the knowledge about daily and seasonal movements and 
population trends of this threatened species (classified as ““vulnerable”’ 
by Collar et al. 1992), we include our observations here. During five 
days spent along the Rio Caine, only one individual was seen (on 
11 June by SH), whereas c. 40 birds were found in the same area on 
1 Aug 1989 (MK pers. obs.), about 60 were estimated to have been 
present in Oct/Nov 1990 (Boussekey et al. 1991) and c. 100 were 
estimated to have been present from Sept 1991 to March 1992 (Pitter & 
Christiansen 1995). This nearly complete lack of sightings of 
Red-fronted Macaws during our survey along the Rio Caine is 
puzzling, especially since the species was not considered to be under 
any immediate threat in this area by Boussekey et al. (1991), and Pitter 
& Christiansen (1995) considered it to be resident. 

At Novillero, MK and SH saw 4 individuals flying downriver at 
dawn on 20 June; none were recorded at this site from 30 April to 
2 May 1995 by P. Nygaard-Andersen and N. Moray Williams (pers. 
comm.). In the Rio Pilcomayo drainage, 22 individuals were seen flying 
to a roost by MK and SH on 22 June about 3 km S of Icla (19°22’S, 
64°48’W). On the evening of the following day between 104 and 116 
birds were observed flying upriver near Hacienda Uyuni (19°26’S, 
64°50°W) by MK and SH, presumably to the same roost as the birds of 
the previous day, whereas only 45 birds flew downriver at the same site 
the next morning. During the day, groups of 3 to 6 birds were found in 
riverine forest along the Rio Pilcomayo (19°27-35’S, 64°49-51'W), 
totalling c. 18 birds along a 20 km stretch of the river. On the morning 
of 7 July about 90 birds were seen flying upriver along Rio La 
Haciendita (19°01’S, 64°12’W) at 1300 m, where none were seen the 
previous afternoon. 


CANARY-WINGED PARAKEET Brotogeris versicolurus 


Observations of a flock of 4-6 individuals by SM on 27 Dec 1993 and 
of several pairs by SKH on 16, 18 and 20 Sept 1995 at Inquisivi 


S. K. Herzog et al. 228 Bull. B.O.C. 1997 117(3) 


represent the first reports of this species for depto. La Paz (Remsen & 
Traylor 1989, Armonia 1995). 


MONTANE FOREST SCREECH-OWL Otus hoyi 

At Rio Azero, this species was captured three times (involving at 
least 2 different individuals) in dry forest at 1100 m whilst mist-netting 
bats on the nights of 28 and 29 June. Additionally, MK heard and saw 
an individual at 1250 m at a site about 1.5 km away from the netting 
area on 27 June. These records fill an altitudinal and ecological gap 
between the humid montane forest site above 1300 m from which the 
species was originally described (K6nig & Straneck 1989) and a recent 
record from the Bolivian chaco (Kratter et al. 1993). At Rio Azero 
O. hoyi is sympatric with O. choliba. 


BUFF-FRONTED OWL Aegolius harnisiu 

The characteristic, wailing trill of this secretive species, for which 
few records exist from Bolivia (J. V. Remsen zn litt.), was heard by MK 
and SH from deciduous forest near our camp at Masicuri on the night 
of 16 July. To our ears the voice was identical to the recording on 
Hardy et al. (1990). The species was also found to be locally common in 
structurally similar, but not mountainous habitat between Pozo 
Colorado and Fn. V. Rivarola, Presidente Hayes, Paraguayan Chaco 
(c. 23°30’S, 60°20’W, 100 m) in Oct 1991 by MK and P. Driesch (cf. 
Paraguayan distribution of this species in Hayes 1995), suggesting that 
it may be more widespread in deciduous forests than previously 
thought, and that the Andean and SE Brazilian populations may be 
connected by a population in the chaco and adjacent dry forest habitats. 


BAND-WINGED NIGHTJAR Caprimulgus longirostris 
A pair seen by MK and SH at 800m on a tributary of the Rio 
Masicuri on the night of 9 July represents a considerable downward 


range extension on the eastern side of the Andes (from 2100 m; Fjeldsa 
& Krabbe 1990, Armonia 1995). 


AMETHYST WOODSTAR Calliphlox amethystina 

Two males were observed by SKH on 7 Oct at 800m along Rio 
Boopi at Las Mercedes. This is, to our knowledge, the first report of 
this species in Andean valleys and represents an increase in altitudinal 
range by 100m (Bond & Meyer de Schauensee 1943, Remsen & 
Traylor 1983). 


STRIPED WOODPECKER Picoides lignarius 
Observations of this species on 25 and 28 Dec 1993 3 by SM and on 
16 Sept 1995 by SKH in dry forest below Inquisivi represent the first 


reports of this species for depto. La Paz (Remsen & Traylor 1989, 
Armonia 1995). 


GREEN-BARRED FLICKER Colaptes melanochloros 


Daily observations of a few individuals between 2200 and 3000 m at 
Inquisivi by SM in 1993 and 1994 and by SKH and SH in 1995 


S. K. Herzog et al. 229 Bull. B.O.C. 1997 117(3) 


represent the first reports for depto. La Paz (Remsen & Traylor 1989, 
Armonia 1995). Interestingly, none of the birds seen in 1995, even 
when two individuals (presumed pairs) were observed together, showed 
the red malar stripe typical of the male. 


BOLIVIAN EARTHCREEPER Upucerthia harterti 

Observations of a pair and a single individual on 13 June by MK and 
SH on the SW side of the Rio Caine represent the first reports for 
depto. Potosi (Remsen & Traylor 1989, Armonia 1995). These birds 
and another pair seen on 24 June at 2500 m in the Rio Jaya Mayu valley 
(17°52'S, 65°56’W) were found in habitat much modified by timber 
extraction, agriculture and grazing. The pair seen along Rio Caine 
spent several hours foraging and vocalizing in a hedge formed by dead 
Acacia branches and a barren cliff, where the birds investigated 
crevices in a manner reminiscent of Rock Wrens Salpinctes obsoletus. 
Remsen ez al. (1988) found this species to be partial to dry scrub and 
forest with a dense undergrowth of terrestrial bromeliads, a typical 
feature of severely overgrazed habitats (MK unpubl. data). These 
observations suggest that U. harterti tolerates human degradation of its 
habitat quite well and may not be “‘near-threatened’’ (as considered by 
Collar et al. 1992). 

An individual of this species was also seen in degraded dry forest 
(dominated by Prosopis laevigata, Caesalpinia spp. and Dodonaea 
viscosa) with abundant terrestrial bromeliads (Puya meziana) at 
Huachillas (16°39’S, 68°01°W, 3000 m) in the La Paz valley on 8 Sept 
1991 by J. Fjeldsa (pers. comm.) and on 9 Sept 1991 by MK. These are 
the first reports of this species for depto. La Paz (Remsen & Traylor 
1989, Armonia 1995). 


Cranioleuca, unnamed species 


A distinctive, rufous-capped form of Cranioleuca spinetail, which 
will be described as a new species by SM and J. Fjeldsa, was a common 
species in dry forest below Inquisivi. It was first discovered and 
tape-recorded by SM on 27 Dec 1993, and specimens were collected a 
few weeks later. The species was mostly seen in pairs in the lower 
canopy and upper understorey but occasionally also in mixed-species 
flocks. It was also frequently encountered by SKH, MK and SH in 
1995. 


BERLEPSCH’S CANASTERO Asthenes berlepschi 


The sighting of a pair at 2300m in the Consata Valley (15°30'S, 
68°38'W) on 30 May by MK and SH represents a new altitudinal 
record for this threatened species (ranked as “insufficiently known’”’ by 
Collar et al. 1992), previously known from 2600-3700 m (Fjeldsa & 
Krabbe 1990), and only recently rediscovered (Mayer 1995). The birds 
were found in a low, open “‘hedge”’ formed by Puya bromeliads along a 
field edge in an area of intense agricultural activity. The Consata Valley 
has been densely inhabited at least since Incan times, and natural 
habitats have been almost completely converted into cultivated areas. 
Nevertheless, A. berlepschi seems to be common within its restricted 


S.K. Herzog et al. 230 Bull. B.O.C. 1997 117(3) 


range (the Consata Valley and its tributaries); three separate individuals 
were seen crossing the road on 3 June by MK while driving from 
Tacacoma to Quiabaya (15°38’S, 68°40'W) at 3200-3500 m in open, 
degraded Polylepis forest and in Baccharis pentlandiu scrub, and the 
species was common around the town of Sorata (Mayer 1995). 
Apparently, A. berlepscht is as common and tolerant of habitat 
degradation within its restricted range as the closely related A. dorbygni 
(of which it may only be a race; Ridgely & Tudor 1994) in similar dry 
valleys further south in Bolivia. 

Our observations increase the known range size of A. berlepschi to 
c. 200 km*. We expect that the species will also be found in the not yet 
surveyed areas north of the Rio Consata where the habitat is very 
similar, which would increase its range size to 450 km’. Based on the 
combined information above, we do not consider A. berlepschi to be 
under any immediate threat and suggest it should be removed from the 
list of threatened species. 


BUFF-BROWED FOLIAGE-GLEANER Syndactyla rufosuperciliata 


Regularly seen by MK and SH in dry forest down to 1000m in 
mixed-species flocks with Poecilurus scutatus, Basileuterus bivittatus and 
Arremon flavirostris at Rio Azero. In the Andes, this species was 
previously known only from humid forest above 1300m (Ridgely & 
Tudor 1994, Armonia 1995). 


SPOT-BILLED GROUND-TYRANT Muscisaxicola maculirostris 


Repeated observations by SKH and MK of an individual along the 
Rio La Paz at 1300 m at Huara represent a downward range extension 
of 700 m for this species in Bolivia (Ridgely & Tudor 1994, Armonia 
1995). 


BROWN-CRESTED FLYCATCHER Myiarchus tyrannulus 


An individual was seen and tape-recorded at 2300m on 15 and 
16 Sept by SKH at Inquisivi. This species was previously known only 
up to about 1700 m (Ridgely & Tudor 1994, Armonia 1995). 


LEMON-BROWED FLYCATCHER Conopias cinchoneti 


One individual of this distinctive flycatcher was observed foraging 
along Rio Khatu below Inquisivi on 16 Sept by SKH. The bird was 
constantly on the move, perching only briefly in two small bushes about 
1-1.5 m above ground on an open gravel bar. It frequently sallied out 
horizontally into the air about 15-25 m away from its perch. The bird 
did not call and disappeared into gallery forest after about 10 minutes. 
On 8 Oct, MK observed another individual along Rio Coroico at 
Yolosillas for about 3 minutes in degraded riverine forest. Both 
individuals were identified by their two-toned colour pattern with olive 
backs and yellow underparts, a yellow superciliary extending far back 
onto the nape and the lack of wingbars and white edgings on the outer 
rectrices. The coloration of the bird at Yolosillas was rather dull, 
suggesting that it was a juvenile. These are the first reports of this 
species for Bolivia and would represent a huge southward range 


S. K. Herzog et al. 231 Bull. B.O.C. 1997 117(3) 
extension of 600km from Cuzco, Peru (Ridgely & Tudor 1994). 


However, until “‘tangible evidence”’ for the occurrence of this species in 
Bolivia is obtained (preferably by specimen), its presence in Bolivia 
should be regarded as hypothetical. 


WHITE-NAPED XENOPSARIS Xenopsaris albinucha 

An individual of this species, known from only a handful of localities 
in Bolivia (J. V. Remsen 77 /itt.), was observed on 16 July by MK ina 
mixed-species flock in tall, disturbed gallery forest along the Rio 
Grande at 500 m in Masicuri. It perched briefly almost directly above 
the observer about 5 m above ground before flying into the viny tangles 
of a treefall gap, where it could not be relocated. 


WHITE-LINED TANAGER Tachyphonus rufus 

A pair seen in degraded riverine forest at 1300 m at Huara on 28 Sept 
by MK represents the first report for depto. La Paz (Remsen & Traylor 
1989, Armonia 1995) and fills part of a large distributional gap for this 
rather local species (Ridgely & Tudor 1989), previously known from 
localities about 750 km further E in eastern Santa Cruz (Armonia 1995) 
and c. 650 km further NW in Cuzco, Peru (Isler & Isler 1987). 


FAWN-BREASTED TANAGER Pipraeidea melanonota 

A pair-seen in a mixed-species flock near the confluence of the Rio 
Masicuri and Rio Grande at 500m on 11 July by MK represents an 
unusually low record of this species on the eastern Andean slope (where 
usually recorded above 1500 m, Ridgely & Tudor 1989; above 1200 m 
according to Armonia 1995). Also regularly seen at 1100-1400 m at Rio 
Azero. 


CHESTNUT-VENTED CONEBILL Conirostrum speciosum 
At Miguillas this species was found at elevations of up to 1500 m by 


SKH, about 500m above the usual altitudinal range of the species 
(Ridgely & Tudor 1989). 


CINEREOUS CONEBILL Conirostrum cinereum 

An individual was observed preening for about 2 minutes in 
degraded riverine forest at 1300 m along the Rio La Paz at Huara on 28 
Sept by MK. This observation represents a considerable downward 


range extension for the eastern Andean slope (previously known only 
above 2500 m; Ridgely & Tudor 1989, Armonia 1995). 


EPAULET ORIOLE I[cterus cayanensis 

A pair was seen at 1500 m at San Juan del Potrero on 5 Sept by 
SKH, 500 m above the usual range for this species (Ridgely & Tudor 
1989, Armonia 1995). 


The following 23 species were found at 2050-2400 m on the SW side 
of the Caine valley (site 7), an area of relatively low elevation, which 
political arbitrariness has placed in depto. Potosi (most of which is well 
above 3000 m), and while they are new to the department (Remsen & 


S.K. Herzog et al. 232 Bull. B:O.C., 4997 1147(3) 


Traylor 1989, Armonia 1995), these records were to be expected based 
on known ranges in adjacent departments and represent only minor 
range extensions: Buff-necked Ibis Theristicus caudatus' (4 on 
12 June by MK and SH), Black-chested Buzzard-eagle Geranoaetus 
melanoleucus (1 on 13 June by MK and SH), Roadside Hawk Buteo 
magnirostris (1 on 11 June by SH), Collared Plover Charadrius collaris 
(2 on 13 June by MK and SH), Large-tailed Dove Leptotila megalura 
(several on 11-13 June by MK and SH), Blue-crowned Parakeet 
Aratinga acuticaudata (common), Smooth-billed Ani Crotophaga ani 
(several on 12 June by MK and SH), Glittering-bellied Emerald 
Chlorostilbon aureoventris (1 male on 12 June by MK), White-bellied 
Hummingbird Amazilia chionogaster (several on 11-13 June by MK 
and SH), Narrow-billed Woodcreeper Lepidocolaptes angustirostris 
(2 on 11 June by MK and SH), Rufous Hornero Furnarius rufus 
(common), Olive-crowned Crescentchest Melanopareia maximiliani 
(1 on 12 June by MK), Suiriri Flycatcher Suirivi suiriri (2 on 13 June 
by MK and SH), Southern Scrub Flycatcher Sublegatus modestus (1 on 
13 June by MK), Greater Wagtail-tyrant Stigmatura budytoides 
(common), Creamy-bellied Thrush Turdus amaurochalinus (several on 
12 June by MK and SH), Grey-crested Finch Lophospingus 
griseocristatus (common), Ringed Warbling-finch Poospiza torquata 
(pair on 13 June by MK and SH), Great Pampa Finch Embernagra 
platensis (singing male by MK and SH on 11 June), Black-backed 
Grosbeak Pheucticus aureoventris (several on 11-13 June by MK and 
SH), Sayaca Tanager Thraupis sayaca (regularly seen by MK and SH), 
Brown-capped Redstart Myioborus brunniceps (2 on 12 June by MK), 
Masked Gnatcatcher Polioptila dumicola (pair on 13 June by MK and 
SH). 


Acknowledgements 


This study was partially funded by the Schimper-Stiftung, Germany. J. Fyjeldsa, 
L. Jammes, J. V. Remsen, F. Sagot and B. Whitney commented on the manuscript. 


References: 

Armonia. 1995. A birdlist of Bolivia. Armonia, Santa Cruz, Bolivia. 

Bond, J. & Meyer de Schauensee, R. 1943. The birds of Bolivia. Part II. Proc. Acad. Nat. 
Sci. Philadelphia 95: 167-221. 

Boussekey, M., Saint-Pie, J. & Morvan, O. 1991. Observations on a population of 
Red-fronted Macaws Ara rubrogenys in the Rio Caine valley, central Bolivia. Bird 
Conserv. Internat. 1: 335-350. 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
T. A. & Wege, D. C. 1992. Threatened birds of the Americas. ICBP/IUCN Red Data 
Book. ICBP, Cambridge, U.K. 

Fjeldsa, J. & Krabbe, N. 1990. Birds of the High Andes. Apollo Books, Svendborg, 
Denmark. 

Hardy, J. W., Coffey, B. B. & Reynard, G. B. 1990. Voices of the New World Owls. ARA 
Records, Gainesville, Florida. 

Hayes, F. E. 1995. Status, Distribution and Biogeography of the Birds of Paraguay. 
American Birding Association. Monographs in Field Ornithology No. 1. 

Hilty, S. L. & Brown, W. L. 1986. A Guide to the Birds of Colombia. Princeton Univ. 
Press. 

Isler, M. L. & Isler, P. R. 1987. The Tanagers. Natural history, distribution and 
identification. Smithsonian Institution Press. 


S. K. Herzog et al. 283 Ball. BOC t997.197(3) 


Konig, C. & Straneck, R. 1989. Eine neue Eule (Aves: Strigidae) aus Nordargentinien. 
Stuttgarter Beitr. Naturk. Ser. A, Nr. 428. 

Kratter, A. W., Sillett, T. S., Chesser, R. T., O’ Neill, J. P., Parker, T. A. & Castillo, A. 
1993. Avifauna of a chaco locality in Bolivia. Wilson Bull. 105: 114-141. 

Mayer, S. 1995. Notes on the occurrence and natural history of Berlepsch’s Canastero 
Asthenes berlepschi. Cotinga 3: 15-16. 

Pitter, E. & Christiansen, M. B. 1995. Ecology, status and conservation of the 
Red-fronted Macaw Ara rubrogenys. Bird Conserv. Internat. 5: 61-78. 

Remsen, J. V., Schmitt, C. G. & Schmitt, D. C. 1988. Natural history notes on some 
poorly known Bolivian birds, part 3. Gerfaut 78: 363-381. 

Remsen, J. V. & Traylor, M. A. 1983. Additions to the avifauna of Bolivia, part 2. Condor 
85: 95-98. 

Remsen, J. V. & Traylor, M. A. 1989. An Annotated List of the Birds of Bolivia. Buteo 
Books, Vermillion, South Dakota. 

Ridgely, R. S. & Tudor, G. 1989. The Birds of South America. Vol. 1. The oscine 
passerines. Oxford Univ. Press. 

Ridgely, R. S. & Tudor, G. 1994. The Birds of South America. Vol. 2. The suboscine 
passerines. Oxford Univ. Press. 


Addresses: S. K. Herzog, Foundation for Tropical Research and Exploration, P.O. Box 
74431, Davis, CA 95617, U.S.A.; current address: Langeooger Str. 8, 26419 
Schortens, Germany. Dr. M. Kessler, Systematisch-Geobotanisches Institut, 
Untere Karspiile 2, 37073 Géttingen, Germany. S. Maijer, Ter Meulenplantsoen 20, 
7524CA Enschede, The Netherlands. S. Hohnwald, Siegstr. 8, 47051 Duisburg, 


Germany. 


© British Ornithologists’ Club 1997 


Appendix 


The following list includes all species observed at 11 of the 12 survey sites described in 
the text (site 12, Rio Pilcomayo, is not included): 1 Consata, 2 Yolosillas, 3 Las Mercedes, 
4 Miguillas, 5 Huara, 6 Inquisivi, 7 Rio Caine, 8 San Juan del Potrero, 9 Novillero, 10 
Masicuri and 11 Rio Azero. The habitat(s) in which each species was observed are 
abbreviated as follows: D=dry forest; E=evergreen forest (including gallery forest); 
A=predominantly agricultural areas; R=directly at rivers; O= flying over the survey area. 
No specimens were collected. Relative abundances were not estimated due to the 
relatively short time spent at each site. 

Tinamus major 10E. Crypturellus obsoletus 3D. Crypturellus undulatus 2DE, 8D. 
Crypturellus atrocapillus 3D, 4D. Crypturellus tataupa 2DE, 3D, 4D, 8D, 10DE. 
Nothoprocta pentlandii 6DA. Phalacrocorax brasilianus 20. Tigrisoma fasciatum 2R. Ardea 
cocot 10R. Ardea alba 10R. Egretta thula 7R, 10R. Nycticorax nycticorax 90. Theristicus 
caudatus 7A. Coragyps atratus 30, 80, 100, 110. Cathartes aura 10, 20, 30, 40, 50, 
60, 70, 80, 90, 100, 110. Vultur gryphus 50, 60, 70, 80, 110. Sarcoramphus papa 
110. Merganetta armata 2R, 6R. Chondrohierax uncinatus 20, 3D, 40, 5D. Elanoides 
forficatus 30. Ictinia plumbea 3D, 40. Accipiter bicolor 90. Geranoaetus melanoleucus 1D, 
30, 50, 60, 70, 80, 90, 110. Harpyhaliaetus solitarius 100, 110. Buteo magnirostris 
1DE, 20, 3D, 4D, 5DE, 6A, 7D, 8D, 9D, 10DE, 11DE. Buteo albigula 60. Buteo 
brachyurus 30, 110. Buteo albicaudatus 80. Buteo polyosoma 90. Buteo albonotataus 10. 
Phalcoboenus megalopterus 60. Polyborus plancus 8D, 11E. Falco sparverius 1D, 3D, 5DE, 
6DE, 7D. Falco femoralis 110. Falco rufigularis 2E, 5D, 10E. Falco peregrinus 50, 70. 
Ortalis guttata 1DE, 2E, 3D, 10D. Penelope montagnii 9D. Penelope jacquacu 3D. Pipile 
pipile 10D. Aramides cajanea 2E, 3E, 10E. Charadrius collaris 7R. Actitis macularia 3R, 
6R. Columba maculosa 70. Columba fasciata 3D, 5D, 6D. Columba plumbea 1D, 3D. 
Zenaida auriculata 1D, 7DA. Columbina talpacoti 10A, 11E. Columbina picui 1D, 3A, 
5DE, 6DA, 7DA, 10DEA. Claravis mondetoura 3D. Leptotila verreauxi 1DE, 2DEA, 3D, 
4D, 5DE, 6DE, 8D, 10DEA, 11DE. Leptotila megalura 3D, 4D, 5DE, 6DE, 7DA, 8D, 
9DE, 11D. Ara militaris 100. Ara rubrogenys 70, 90. Ara auricollis 10E, 11DE. Ara 
severa 100. Aratinga acuticaudata 7DA, 8D. Aratinga mitrata 1D, 20, 3D, 4D, 5DE, 
6D, 7D, 9DE, 11DE. Pyrrhura molinae 2DE, 3D, 4D, 5DE, 6DE, 10DE, 11D. 
Myiopsitta monachus 7DA. Bolborhynchus aymara 6D, 7D. Brotogeris versicolurus 1D, 


S. K. Herzog et al. 234 Bull. B.O.C. 1997 117(3) 


6DE, 70, 90, 100, 110. Pionus menstruus 10, 20, 3D. Pionus maximiliani 8D, 100, 
110. Pionus sp. 40. Amazona aestiva 70, 8D, 100, 110. Piaya cayana 1D, 2DE, 3D, 
4D, 5DE, 6D, 8D, 10E, 11DE. Crotophaga ani 7A, 10A. Otus choliba 4D, 6DA, 8D, 9D, 
10E, 11D. Otus hoyi 11D. Bubo virginianus 60. Pulsatrix perspicillata 3D, 10E. 
Glaucidium brasilianum 10D. Glaucidium sp. 2E. Aegolius harrisii 10D. Nyctibius griseus 
4D. Nyctidromus albicollis 11D. Caprimulgus rufus 4D, 5D. Caprimulgus longirostris 9DE, 
10ER. Hydropsalis brasiliana 8D. Uropsalis lyra 60. Cypseloides rutilus 60. Streptoprocne 
zonaris 60, 80. Aeronautes montivagus 50. Aeronautes andecolus 60, 70, 90. Panyptila 
cayennensts 100. Phaethornis superciliosus 10E. Phaethornis pretret 8D, 11D. Campylop- 
terus largipennis 2E. Colibri thalassinus 2DE. Colibri serrirostris 6D. Chlorostilbon 
mellisugus 1DE, 3D, 4D, 5DE, 6DE. Chlorostilbon aureoventris 7DA, 8D, 9D, 11D. 
Thalurania furcata 10E. Taphrospilus hypostictus 10E. Amazilia chionogaster 1D, 2DE, 
3D, 4D, 5DE, 6DEA, 7A, 8D, 9DE, 10DE, 11DE. Oreotrochilus adela 7D. Patagona 
gigas 7D. Sappho sparganura 5D, 7D, 9D, 11D. Heliomaster longirostris 2D. Calliphlox 
amethystina 3D. Acestrura mulsant 2D. Trogon collaris 10E. Trogon curucui 2E, 3D, 4D, 
5E. Electron platyrhynchum 3D. Momotus momota 3D, 4D, 5DE, 8D, 10DE, 11DE. 
Ceryle torquata 2ER, 10ER. Chloroceryle amazona 2ER. Chloroceryle americana 10ER, 
11ER. Nystalus chacuru 1D, 6A, 10D. Nystalus maculatus 8D. Monasa mgrifrons 10E. 
Pteroglossus castanotis 3D. Ramphastos tucanus 3D. Ramphastos toco 10E, 11E. Picumnus 
cirratus 3D, 5D, 8D, 10D, 11D. Picoides lignarius 6D, 7D, 9D. Veniliornis fumigatus 4D, 
5E. Veniliornis frontalis 5D, 11D. Veniliornis sp. 10E. Piculus chrysochloros 10E. Piculus 
rubiginosus 2E, 8D, 11D. Colaptes melanochloros 6DA, 9D. Dryocopus lineatus 3D, 4D, 
5D. Campephilus melanoleucos i0DE, 11D. Campephilus rubricollis 4D. Campephilus 
leucopogon 8D, 9D. Sittasomus griseicapillus 3D, 4D, 5D, 8D, 10DE, 11D. Xiphocolaptes 
promeropirhynchus 3D, 10E. Xiphocolaptes major 10D, 11D. Dendrocolapies picumnus 4D, 
8D. Xiphorhynchus guttatus 10E. Lepidocolaptes angustirostris 7D, 8D, 9D, 11D. 
Lepidocolaptes albolineatus 3D. Campylorhamphus trochilirostris 4D, 5D. Upucerthia 
hartertt 7DA, 8D. Cinclodes fuscus 5R, 7R, 9R, 11R. Furnarius rufus 6A, 7A, 8D, 9A, 
10A. Synallaxis azarae 1DE. Synallaxis frontalis 5E, 6DEA, 8D, 10DE. Poecilurus 
scutatus 3D, 10E, 11D. Cranioleuca pyrrhophia 8D, 9DE. Cranioleuca, unnamed species 
6DEA. Asthenes dorbignyi 6DA. Asthenes berlepschi 1A. Phacellodomus striaticeps 6A, 8D. 
Syndactyla rufosuperciliata 11D. Phylidor rufus 10E. Xenops rutilans 1ODE, 11D. Batara 
cinerea 11E. Taraba major 3D, 4D, 5DE, 10DE. Thamnophilus doliatus 2E, 3D, 4D, 5EA. 
Thamnophilus aroyae 2E. Thamnophilus punctatus 1ODE. Thamnophilus caerulescens 1E, 
5DE, 6DE, 8D, 9DE, 11D. Thamnophilus ruficapillus 11D. Thamnophilus sp. 4D. 
Myrmotherula longicauda 2E. Herpsilochmus atricapillus 10E. Formicivora melanogaster 
10E. Pyriglena leuconota 3D, 4D, 6E. Chamaeza campanisona 11E. Melanopareia 
maximiliani 5D, 6A, 7D. Phyllomyias burmeisteri 5E. Phyllomyias sclatert 10E. Zimmerius 
bolivianus 4D. Camptostoma obsoletum 3D, 4D, 5D, 6DEA, 8D, 11D. Phaeomyias murina 
6A, 8D. Sublegatus modestus 5D, 7D, 10D. Suiriri suiriri 7D, 9D, 10E. Myiopagis 
gaimardu 3D. Myiopagis viridicata 1E, 10E. Elaenia obscura 6EA. Elaenia sp. 3D, 8D. 
Mecocerculus leucophrys 9D. Serpophaga cinerea 2R, 3R. Serpophaga munda 5D, 6DE, 
7D, 8D, 11D. Stigmatura budytoides 7DA, 8D, 9D. Anairetes flavirostris 6D, 7DA. 
Leptopogon amaurocephalus 2E, 3D, 10E. Leptopogon superciliaris 3D. Phylloscartes 
ophthalmicus 2E, 3D. Phylloscartes ventralis 11D. Hemitriccus margaritaceiventer 1D, 2D, 
3D, 4D, 5D, 6D, 8D, 10DE, 11D. Todirostrum cinereum 10E. Tolmomyias sulphurescens 
2E, 3D, 4D, 10E. Myiophobus fasciatus 2E, 5E, 6 DEA. Pyrrhomyias cinnamomea 6E. 
Contopus fumigatus 5E, 6E. Contopus cinereus 10E. Empidonax sp. 3D. Lathrotriccus euleri 
3D. Cnemotriccus fuscatus 5D. Sayornis nigricans 1R, 2R, 3R, 5R,. 6ER, 10R, 11R. 
Pyrocephalus rubinus 3D. Ochthoeca leucophrys 9D. Muscisaxicola maculirostris 5R, 7R. 
Knipolegus aterrimus 3D, 4D, 5DE, 6A, 7DA, 8D, 9DE, 11DE. Colonia colonus 10E. 
Satrapa icterophrys 3D. Hirundinea ferruginea 3D, 5D, 6DE, 7DA, 8D, 9DE, 10D. 
Machetornis rixosus 10A. Casiornis rufa 2E, 3D, 4D, 5D, 10E. Myiarchus swainsoni 11D. 
Myiarchus ferox 3D. Myiarchus cephalotes 2DE. Myiarchus tyrannulus 3D, 4D, 5DE, 6D, 
11D. Myiarchus sp. 8D, 10E. Pitangus sulphuratus 7A, 9DEA, 10EA, 11DE. Mytozetetes 
similis 2E. Conopias cinchoneti 2E, 6ER. Myiodynastes chrysocephalus 3D. Mytiodynastes 
maculatus 2E, 3D, 4D, 5D. Legatus leucophaius 2E. Tyrannus melancholicus 1DE, 2E, 3D, 
SDE, 6EA, 10EA, 11DE. Xenopsaris albinucha 10E. Pachyramphus viridis 10E. 
Pachyramphus castaneus 3D. Pachyramphus polychopterus 3D. Phytotoma rutila 7DA, 9A. 
Notiochelidon cyanoleuca 2E, 3D, 5E, 7A, 9AO, 10E. Atticora fasciata 3DR. Thryothorus 
genibarbis 1DE, 2DE, 10DE. Troglodytes aedon 2DEA, 3DA, 4D, 5DEA, 6DE, 7DA, 8D, 
9DEA, 11DE. Cinclus leucocephalus 1R. Polioptila dumicola 7DA, 8D, 9D, 11D. 


In Brief 235 Bull. B.O.C. 1997 117(3) 


Myadestes ralloides 2E. Entomodestes leucotis 5E. Turdus chiguanco 6A, 7A, 9EA, 11D. 
Turdus fuscater 6A. Turdus rufiventris 8D, 1ODE, 11DE. Turdus amaurochalinus 1D, 2D, 
SDE, 6DE, 7DA, 10DE. Mimus dorsalis 7DA. Cyanocorax cyanomelas 2DE, 3D, 4D, 
5DE, 8D, 10DE, 11DE. Cyanocorax chrysops 8D, 10DE, 11D. Cyclarhis gujanensis 4D, 
5D, 6D, 8D, 10DE. Vireo olivaceus 1D, 2E, 3D, SDE, 6DA, 10DE. Hylophilus 
hypoxanthus 10E. Zonotrichia capensis 4D, 7A, 8D, 9EA. Ammodramus aurifrons 3RA, 
5R, 10A. Lophospingus griseocristatus 7DA, 8D, 9DA. Poospiza boliviana 9D. Poospiza 
whiti 6DE. Poospiza torquata 5DE, 6DE, 7D, 8D, 9D. Poospiza melanoleuca 8D, 11D. 
Sicalis flaveola 6A, 8D. Embernagra platensis 7A, 10DE. Volatinia jacarina 2A. 
Sporophila caerulescens 3D, 5DE, 6A. Catamenia analis 5E, 6DA. Tiaris obscura 3D, 
5DE. Arremon flavirostris 4D, 5DE, 8D, 9D, 10D, 11D. Atlapetes fulviceps 9E. Aitlapetes 
torquatus 9E. Coryphospingus cucullatus 1D, 2EA, 3DA, 4D, 5DE, 8D, 10D, 11D. 
Pheucticus aureoventris 4D, 5DE, 6DE, 7A, 8D, 9EA, 10E. Saltator maximus 1DE, 2DE. 
Saltator aurantirostris 6DE, 7A, 8D, 9DEA. Cyanocompsa brissoni 2E, 4D, 5E, 8D. 
Schistochlamys melanopis 1E. Cissopis leveriana 10E. Chlorospingus ophthalmicus 6D. 
Thlypopsis sordida 10E. Hemithraupis guira 4D, 5D, 10E. Tachyphonus rufus 5E. Piranga 
flava 2E, 5D, 8D, 9DE, 10DE, 11D. Ramphocelus carbo 1DE, 2E, 3DA, 10E. Thraupis 
sayaca 1DE, 3DA, 4D, 5DE, 6DE, 7A, 8D, 9DE, 10DE, 11DE. Thraupis palmarum 1E, 
3D. Thraupis bonariensis 5E, 6DE, 7A, 8D. Pipraeidea melanonota 10E, 11D. Euphonia 
chlorotica 8D. Euphonia laniirostris 2E, 3D. Euphonia cyanocephala 3D, 4D, 5D, 6DE. 
Euphonia sp. 2E. Cyanerpes caeruleus 1E. Diglossa sittoides 2E, 4D. Tersina viridis 2E. 
Coereba flaveola 2DE, 3D, 4D, 5DE, 8D. Parula pitiayumi 2DE, 4D, 5DE, 6DE, 8D, 
10DE, 11DE. Geothlypis aequinoctialis 2E, 1OE. Myioborus brunniceps 6DE, 7A, 8D, 9E, 
11DE. Basileuterus bivittatus 1D, 2DE, 3D, 5D, 9DE, 10DE, 11D. Basileuterus coronatus 
2E. Conirostrum speciosum 3D, 4D, 5D. Conirostrum cinereum 5E. Psarocolius decumanus 
1DE, 2DE, 3D, 4D, 5DE, 10DE, 11DE. Psarocolius atrovirens 2E, 3D, 5DE, 6D. 
Psarocolius angustifrons 3D. Cacicus chrysopterus 11D. Icterus cayanensis 8D, 10E, 11D. 
Oreopsar bolivianus 7A. Molothrus badius 6A, 7A, 9A, 10A. Scaphidura oryzivora 3DA. 
Carduelis magellanica 7A, 9E. Carduelis xanthogastra 6A. Carduelis sp. 2E, 3D. 


IN BRIEF 


ON THE IDENTITY OF LOPHORNIS MELANIAE FLOERICKE 
(TROCHILIDAE) 


In 1920, Curt Floericke described a new species of Lophornis, on the 
basis of two unlocalised males. He described his new species, L. 
melaniae, as intermediate between L. stictolopha and L. delattret. 
According to Floericke, the crest feathers were narrow and pointed but 
not nearly so “‘radical or thread-like’’ as those of delattrez, and they all, 
or nearly all, had a black terminal spot but this was much smaller than 
in stictolopha. Floericke admitted that melaniae looked superficially like 
aberrant specimens or hybrids between the other species, but this he 
considered unlikely, as he recalled seeing other similar specimens 
before, and furthermore the crest was not intermediate in colour, but 
even paler than in delattrei, especially in the central part which pales to 
a pale isabelline yellow. The bill was stated to be longer than in either 
stictolopha or delattret. 

Lophornis delattrei occurs through tropical and subtropical zones in 
southwest Mexico and from the Pacific slopes of Costa Rica and 
Panama through the central and eastern Andes of Colombia to northern 
and eastern Peru and Bolivia. The male has a rather long chestnut-buff 
crest, each feather sharply pointed and tipped with a tiny green 
“spangle’’. The cheek feathers are only slightly lengthened, rufous with 
green tips. The monotypic species Lophornis stictolopha occurs through 


In Brief 236 Bull. B.O.C. 1997.117(3) — 


western Venezuela, eastern Colombia and eastern Ecuador. The male is 
very like that of delattrez but the crest feathers are not narrowed to a 
point, and are tipped with larger spots, black but shining faintly green 
from some angles and mauve from others. 

There are three (or perhaps four) recognised races of delattrei, 
including brachylopha from Mexico, and lessoni from Costa Rica to 
Colombia, in which the male differs from the nominate in that the 
points of the crest feathers are sharper, and the spangles in some 
specimens are almost or quite absent. Nominate delattrei is from 
northeastern and central Peru. A fourth race, regulus, from Bolivia, is 
questionably distinct from delattrez; Zimmer (1950) could find no 
difference in adult males but some possible differences in females. 

It has been suggested by Meyer de Schauensee (1966) that melaniae 
represented melanistic specimens of some other species of Lophornis, 
but he appears to have based this assumption on the name, since he did 
not see either the specimens or the original reference. Neither the name 
nor the description has anything to do with melanism: Floericke named 
the bird after his wife! (Meyer de Schauensee’s English name “‘Dusky 
Coquette”’ is therefore also inappropriate.) 

It is not possible to assess the types, for these almost certainly no 
longer exist. I am assured by Dr H-W. Mittmann (2m litt.) that 
Floericke’s collection was stored at the Naturalienkabinett Stuttgart, 
and was completely destroyed during World War Two. All discussion 
must therefore be somewhat subjective. However, while curating the 
Natural History Museum collection, I found a pale-crested skin with 
reduced spangling, which agreed quite closely with Floericke’s 
description of melaniae. Males of L. delattrei exhibit variation in depth 
of colour of the crest, so Floericke may have had some abnormally pale 
(or perhaps faded) specimens. Thus, Lophornis melaniae has no 
taxonomic validity, and probably represents aberrant or faded 
specimens of L. delattrez. 


I am grateful to Derek Goodwin for kindly translating Floericke’s paper, and to Gary 
Graves for commenting on an early draft of this note. 


References: 

Floericke, C. 1920. Neue Kolibriformen, Mitt. Vogelwelt 19: 2-4. 

Meyer de Schauensee, R. 1966. The Species of Birds of South America and their 
Distribution. Academy of Natural Sciences, Philadelphia. 

Zimmer, J. T. 1950. Studies of Peruvian birds no. 57. Amer. Mus. Novit. 1463. 


The Natural History Museum, MICHAEL WALTERS 
Akeman St., Tring, 
Hertfordshire HP23 6AP, U.K. 15 July 1996 


In Brief 237 Bull. B.O.C. 1997 117(3) 
THE MALEO MACROCEPHALON MALEO ON BUTON 


The Maleo Macrocephalon maleo is an endemic megapode of Sulawesi, 
Indonesia (White & Bruce 1986, Jones et al. 1995). Its distribution and 
the status of nesting grounds on the island have been studied in detail 
(Dekker 1990, Argeloo 1994). However, the occurrence of the species 
on Sulawesi’s off-lying islands is obscure and requires confirmation 
(Argeloo 1994, Jones et al. 1995). 

Buton, one of the islands where according to indirect evidence 
maleos could possibly occur (Dekker 1990, Argeloo 1994), is located 
to the south of Sulawesi Tenggara, the southeastern province. Until 
recently, there had been no confirmed information regarding the 
occurrence of Maleos on Buton other than eggs which were for sale at 
the market of Maligano and which were said to originate from a 
sandy area around the headwaters of the Lebo river (Pramono 1991). 
Early ornithological expeditions did not mention the species for the 
island (see van Bemmel & Voous 1951). Recently, Addin (1992) 
studied microhabitat characteristics of the Maleo nesting grounds and 
reported the presence of Maleos along the Lebo and Lagito rivers 
(North Buton Wildlife Reserve between 122°47' and 123°13’ E). 
Sykes (1996) reported their presence on the Maligano coast (between 
4°20’ and 5°38’ S). An average of 4.5 Maleo pairs visited the nesting 
ground at the Lebo river per day between July and September 1991, 
with a minimum of 3 pairs and a maximum of 6 pairs per day (Addin 
1992). Approximately 10 birds were recorded on the Maligano coast 
during a three-month survey between August and November 1995 
(Sykes 1996). These data confirm the occurrence of the species on 
Buton. 


References: 

Addin, A. 1992. Characteristics of micro habitat of Maleo (Macrocephalon maleo 
Sal. Muller 1946) in nature in relation to breeding effort at North Buton Nature 
Reserve, South East Sulawesi. Unpublished Sarjana thesis, Bogor Agriculture 
Univ. 

Argeloo, M. 1994. The Maleo Macrocephalon maleo: new information on the distri- 
bution and status of Sulawesi’s endemic megapode. Bird Conserv. Internat. 4: 
383-393. 

Dekker, R. W. R. J. 1990. The distribution and status of nesting grounds of the 
Maleo Macrocephalon maleo in Sulawesi, Indonesia. Biological Conservation 51: 
139-150. 

Jones, D. N., Dekker, R. W. R. J. & Roselaar, C. S. 1995. The Megapodes. Oxford Univ. 
Press. 

Pramono, A. H. 1991. Maleo on Buton. Kukila 5: 150. 

Sykes, B. R. (ed.) 1996. Results of the 1995 Bird Survey of Buton Island South East 
Sulawesi and proposed work programme for 1996. Report for Operation Wallacea. 
Ecosurveys Ltd., Lincolnshire. 

Van Bemmel, A. C. V. & Voous, K. H. 1951. On the birds of the islands of Muna and 
Buton, SE Celebes. Treubia 21: 27-104. 

White, C. M. N. & Bruce, M. D. 1986. The Birds of Wallacea (Sulawesi, the Moluccas and 
Lesser Sunda islands, Indonesia). B.O.U. Checklist No. 7. 


Balitbang Zoologi, DEWI M. PRAWIRADILAGA 
P.O. Box 230, 


Boger 16004, Indonesia. 14 October 1996 


In Brief . 238 Bull. B.O.C. 1997 117(3) 
FRINGILLA COELEBS GENGLERI—AUTHORSHIP AND DATE REVISITED 


Some time ago, in a note in this journal, Clancey (1993) stated that the 
British race of the Chaffinch, known as Fringilla coelebs gengleri 
Kleinschmidt (1909), should instead be attributed to Hens & van Marle 
(1933). According to Clancey, Kleinschmidt’s description of gengleri as 
‘“‘form. nov.” introduced an unavailable infrasubspecific entity, and 
Hens & van Marle were the first to use the name for a subspecies, thus 
making it available under Art. 10 (c) of the Code (International 
Commission on Zoological Nomenclature 1985) and becoming its 
authors following Arts. 50 (c) (i) and 23 (j). This view is mistaken and 
obviously originated from certain misinterpretations of the rules and 
the original work. It seems worthwhile to use this case to illustrate the 
provisions of the Code regulating treatment of infraspecific names 
which are rather complicated and distributed among various articles. 

First of all, if the interpretation were actually correct that 
Kleinschmidt (1909) had created with F. c. gengleri an infrasub- 
specific name in the sense of the Code, it would not have been Hens & 
- van Marle (1933) who made it available, but most probably Gengler 
(1924). In this detailed study of Chaffinch races, Fringilla coelebs 
genglert Kleinschmidt appears among others, thus Art. 10 (c) is 
satisfied. It cannot be excluded, however, that some other work 
between Kleinschmidt’s and Gengler’s publications already uses 
genglert as a name for a subspecies. I only mention these details to 
highlight an awkward nomenclatorial problem when author and date of 
names first published as infrasubspecific have to be determined: 
complete knowledge of pertinent literature is necessary. 

Fortunately, in the case discussed here, this problem is not relevant. 
Clancey’s allegation that F. c. genglert is not available from 
Kleinschmidt (1909) is obviously based on the belief that an intersexual 
variant was described as a “form. nov.”, being expressly infrasub- 
specific under Art. 45 (f) (iv). A study of the original work reveals, 
however, that this is not the case. Kleinschmidt first describes an 
individual variant in Chaffinches, then states that “*... I found among 
English Chaffinches such a surprisingly high percentage of ... (this 
variant) ... that I dare separate the English breeders because of the 
frequency of the new variant’’. It is thus clear that the name gengleri 
was erected not for the individual variant itself but for a geographic 
entity. Kleinschmidt himself, in the very same paper, says that “‘.. . it 
is debatable if such an individual variety should be named as an aberr. 
nov. as insect collectors do in similar cases’’. Throughout his scientific 
work, Kleinschmidt used ‘‘form’’ as a denomination for geographic 
subdivisions within his ‘‘Formenkreise’’, a kind of precursor of the 
superspecies concept (e.g. Kleinschmidt 1900, 1901). While the status 
of Kleinschmidt’s “‘Formenkreis’’-names is debatable because of his 
deliberate disregard of the rules of zoological nomenclature, all of his 
names erected for “‘forms’’ doubtlessly have to be treated as available 
names in the species-group following Arts. 45 (f) (ii) and (g) (11) (for an 
overview of Kleinschmidt’s nomenclature, see Haffer (1992) and 
references therein). 


In Brief 239 Bull. B.O.C. 1997 117(3) 


Finally, another tricky detail of the regulations of the ICZN can be 
illustrated here. For a name published expressly for a “‘form’’, Art. 45 
(g) is to apply. Thus, even if gengleri had indeed been published 
explicitly for an individual variant, it would not be deemed 
infrasubspecific, because it was adopted for a subspecies prior to 1985 
(namely by Gengler (1924) as stated above) and would thus retain its 
original author and date [Art. 45 (g) (ai) (1)]! This does not hold, 
though, for the fourth names in quadrinominal combinations which are 
always infrasubspecific under Art. 45 (f) (111). 

To summarize, no interpretation of the rules is possible that could 
justify Clancey’s assumption that Kleinschmidt’s name was infrasub- 
specific. Therefore, if one indeed wishes to separate the British 
Chaffinches at subspecies level, the name must clearly be Fringilla 
coelebs gengleri Kleinschmidt (1909). 


I am indebted to Dr Ward Tomlinson for linguistical improvements to the manuscript. 


References: 

Clancey, P. A. 1993. The authorship of the name of the British race of the Chaffinch. 
Bull. Brit. Orn. Cl. 113: 189-190. 

Gengler, J. 1924. Buchfinken-Studien. Verh. Orn. Ges. Bayern 16: 103-128. 

Haffer, J. 1992. The history of species concepts and species limits in ornithology. Bull. 
Brit. Orn. Cl. 112A: 107-158. 
Hens, P. A. & van Marle, J. G. 1933. Aanteekeningen omtrent enkele vermeende 
subspecies van den vink, Fringilla coelebs L. Org. Club Nederl. Vogelk. 6: 49-58. 
International Commission on Zoological Nomenclature. 1985. International Code of 
Zoological Nomenclature, 3rd edition. International Trust for Zoological Nomencla- 
ture, London. 

Kleinschmidt, O. 1900. Arten oder Formenkreise? J. Orn. 48: 134-139. 

Kleinschmidt, O. 1901. Der Formenkreis Falco Hierofalco und die Stellung des 
ungarischen Wirgfalken in demselben. Aquila 8: 1-49. 

Kleinschmidt, O. 1909. Fringilla caelebs gengleri, form. nov. Falco 5: 13. 


Institut fir Zoologie, ROBERT GUSTEN 
Technische Hochschule Darmstadt, 

Schnittspahnstr. 3, 

64287 Darmstadt, Germany. 1 August 1996 


MELANISM IN THE GULLS (LARIDAE) 


Melanism in gulls is an uncommon phenomenon. In my previous 
review (Sage 1963) I was aware of just eleven records involving three 
species—Black-headed Gull Larus ridibundus, Herring Gull L. 
argentatus and Lesser Black-backed Gull L. fuscus. 'There is always the 
possibility of some examples of melanism reported in gulls being due to 
oiling or some other form of soiling, rather than pigmentation. 
However, the melanism of a Herring Gull reported by Stokoe (1954) 
was almost certainly genetically based (see Sage 1963), and in the case 
of two melanistic Black-headed Gulls seen in Northumberland in the 
winter of 1961-62, it was established that in the case of at least one bird 


In Brief 240 Bull. B.O.C. 1997 117(3) 


the condition was due to pigmentation and not soiling (Sage 1962). It is 
of interest that Frantzen (1963) described a melanistic Black-headed 
Gull seen in Germany two months before the Northumberland record. 
An individual of this species showing both albinism and melanism 
which appeared to be genetically based was described by Harrison & 
Harrison (1962). 

Since my earlier paper, three further records of melanism in the 
Black-headed Gull have come to hand and it is important that they be 
published in order to complete the record. One with the head, back and 
wings black; tail and underparts grey; legs reddish; seen at Chew Valley 
Lake, Somerset, on 23 January 1966 (A. P. Radford in litt.). One a 
uniform sooty-brown all over, bill and leg colour not seen, was present 
at Bosterne, Hampshire, on 15 March 1967 (Dr J. S. Ash zm litt.). One 
with the head and wings deep charcoal-grey, back and underparts 
black, seen at Lowestoft Ness, Suffolk, on 15 November 1970 (W. H. 
Jolly zn litt.). 

A photograph and description of a melanistic Black-headed Gull seen 
on Sanday, Orkney, in July 1996 has been published recently by 
Thorne (1996). Finally, I was previously unaware of the record of an 
almost completely melanistic Laughing Gull L. atricilla seen in 


Florida, U.S.A., in October 1932 (Westen 1934). 


References: 

Frantzen, M. 1963. Eine melanistische Lachmowe (Larus ridibundus) bei Braunschweig. 
JF. Orn. 104: 252-253. 

Harrison, J. M. & Harrison, J. G. 1962. A Black-headed Gull showing both albinism and 
melanism. Brit. Birds 55: 435-436. 

Sage, B. L. 1962. Albinism and melanism in birds. Brit. Birds 55: 201-225. 

Sage, B. L. 1963. The incidence of albinism and melanism in British birds. Brit. Birds 
56: 409-416. 

Stokoe, R. 1954. A black Herring Gull in Cumberland. Brit. Birds 47: 132-133. 

Thorne, R. H. F. 1996. Melanistic Black-headed Gull. Brit. Birds 89: 570. 

Westen, F. M. 1934. A melanistic Laughing Gull at Pensacola, Florida. Auk 51: 82-83. 


Waveney House, Waveney Close, BRYAN SAGE 
Wells-next-the-sea, 
Norfolk NR23 1HU, U.K. 7 December 1996 


. NOTICE TO CONTRIBUTORS 

Papers, from Club Members or non-members, should be sent to the Editor, Dr 
C. J. Feare, 2 North View Cottages, Grayswood Common, Haslemere, Surrey 
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Registered Charity No. 279583 


CONTENTS 


CLUB NOTICES. Annual General Meeting. Meetings. Accounts........ 
MADRONO N., A. & ESQUIVEL, E. Z. Noteworthy records and range 
extensions from the Reserva Natural del Bosque Mbarayacu 
(Mbaracayu Forest Nature Reserve), Departamento de Canindeyu, 
Paraguay o. 04 6 oes oak a Sk Rk CS ee er 
ALSTROM, P., OLSSON, U. & COLSTON, P. R. Re-evaluation of the 
taxonomic status of Phylloscopus proregulus kansuensis Meise...... 
BARROWCLOUGH, G., LENTINO R., M. & SWEET, P. R. New records of birds 
from Auyan-tepui, Estado Bolivar, Venezuela ................. 
FRITH, C. B. & FRITH, D. Ww. A distinctive new subspecies of Macgregor’s 
Bowerbird (Ptilonorhynchidae) of New Guinea................ 
LOUSADA, S. A. & HOWELL, S. N. G. Amazona oratrix hondurensis: a new 
subspecies of parrot from the Sula Valley of northern Honduras .. 
DUCKWORTH, J. W. Observations on a population of Jerdon’s Bushchat 
Saxicola jerdoni in the Mekong channel, Laos................. 
ALEIXO, A. Range extension of the Large-headed Flatbill Ramphotrigon 
megacephala with comments on its distribution ................ 
HERZOG, S. K., KESSLER, M., MAIJER, S. & HOHNWALD, S. Distributional 
notes on birds of Andean dry valleys in Bolivia................ 
In Brief WALTERS, M. On the identity of Lophornis melaniae Floericke 


(Trochilidae) . 2.006 6.00. «oe ed Be er 
PRAWIRADILAGA, D. M. ‘The Maleo Macrocephalon maleo on 
Buton 2. ce bie ds a We ake wie eee arr 
GUSTEN, R. Fringilla coelebs gengleri—authorship and date 
revisited 3.22.2. 0). Gawd a oo wie ee 


SAGE, B. Melanism in the gulls (Laridae)................ 


238 
239 


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mail, after their current subscription has been paid. 


COMMITTEE 
Revd T. W. Gladwin (Chairman) (1997) Miss H. Baker (1994) 
Mrs A. M. Moore (Vice-Chairman) (1997) Dr R. P. Prys-Jones (1995) 
Cdr M. B. Casement, OBE, RN N. H. F. Stone (1995) 
(Hon. Secretary) (1996) D. Griffin (1997) 
D. J. Montier (Hon. Treasurer) (1997) N. J. Redman (1997) 


Hon. Editor: Dr D. W. Snow—Dr C. J. Feare, as from September 1997 
Chairman of Publications Sub-committee: Mrs A. M. Moore 
Publications Officer: S. J. Farnsworth 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


QOL 
61 ISSN 0007-1595 
B%62 
Bi RO 
Bulletin of the 


British Ornithologists’ Club 


Edited by 
Dr D. W. SNOW and Prof C. J. FEARE 


JAN 2 5] O98 


LIBRARIES 


Volume 117 No. 4 December 1997 


FORTHCOMING MEETINGS 


Advance notice of meeting dates for 1998. Nine meetings are being arranged for the 
following Tuesdays: 20 January (Ian Burrows on Birds of Papua New Guinea—see below), 
17 March (Tony Prater on Waders), 21 April (Dr W. R. P. (Bill) Bourne on Birds and Islands), 
19 May (AGM and social evening—with informal “mini-talks’” as in 1997), 14 July, 
15 September, 13 October and 17 November. Details of speakers on these dates will be 
published when finalised. 


Tuesday 20 January 1998. Dr Ian Burrows B.Sc., Ph.D. will speak on “The Birds of 
Papua New Guinea’’. Ian has been a keen birder for many years, with a passion for rare and 
elusive species. After spending a year as a Warden of Cape Clear Bird Observatory in 1975, he 
obtained a Ph.D. in Applied Microbiology from Aston University in 1980. He has spent ten 
years (1986-1995) in Papua New Guinea lecturing in Microbiology, Ecology and Biology at the 
University of Papua New Guinea, in Port Moresby. He has visited and watched birds in every 
province of PNG, apart from Bougainville, and has conducted extensive research and 
consultancy programmes on the Melanesian Scrubfowl, Macgregor’s Bird of Paradise, and a 
wide range of rainforest birds. A fluent pidgin speaker, he has worked as an in-field consultant 
for the British and Australian Broadcasting Corporations and the World Wildlife Fund. In 
1997, together with Phil Gregory, he has founded Sicklebill Safaris Ltd., specialising in tours 
to Melanesia and Australia in search of rare and little known species. 

Applications to the Hon. Secretary by 4 Fanuary, please. 


Tuesday 17 March 1998. Dr A. J. (Tony) Prater, Ph.D, B.Sc will speak on ““Waders”’. 
Having obtained his BSc in zoology (Exeter, 1965), Tony has been actively involved with birds, 
travelling widely, especially to North and South America, Africa, Europe, Australia and the 
Antarctic. His first appointment was as Warden of Lundy Bird Observatory in 1965. Since then 
he has held many posts with the RSPB and BTO. His special interest in waders began whilst 
working for the RSPB, on wader feeding as part of the feasibility study for the Morecambe Bay 
Barrage (1968-70). He is the author of several books, including Identification and ageing of 
Holarctic Waders (1977), Estuary Birds of Britain and Ireland (1981), and Shorebirds (1986). 
Since 1994 he has been Deputy RSPB Officer for Wales. 

Applications to the Hon. Secretary by 3 March, please. 


Tuesday 21 April 1998. Dr W. R. P. (Bill) Bourne will speak on “Birds and Islands”. At 
the age of ten, Bill was evacuated to Bermuda, in the central western North Atlantic, for three 
years during World War II, where he sailed and fished and chased the local birds. Since then, 
in the course of a complicated career ending with an Honorary Research Fellowship at 
Aberdeen University, he has periodically managed to spend shorter periods watching both the 
land and seabirds of a variety of other islands, ranging from Bear Island and Cyprus, via the 
Cape Verde Islands and Ascension, to South Georgia and Juan Fernandez. He will summarise 
these experiences and draw some conclusions. 

Applications to the Hon. Secretary by 7 April, please. 


Meetings are held in the Sherfield Building of Imperial College, South Kensington, London 
SW7, at 6.15 p.m. for 7 p.m. The nearest Tube station is at South Kensington, and car parking 
facilities are available; a map of the area will be sent to members, on request. The cash bar is open 
from 6.15, and a buffet supper, of two courses followed by coffee, is served from about 7.00. (A 
vegetarian menu can be arranged if ordered at the time of booking.) Informal talks are given on 
completion. 


Overseas Members visiting Britain are particularly welcome at meetings. For 
details in advance, please contact the Hon. Secretary, Cdr M. B. Casement, OBE, RN: 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1997 


Apart from single copies made for the purposes of research or private study, or criticism or 
review, as permitted under UK law, no part of this publication may be reproduced, stored 
or transmitted in any form or by any means, except with prior permission in writing of the 
publishers, or in accordance with the terms of licences issued by the Copyright Licensing 
Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; for 
address see inside back cover. 


a 


— 


241 Bull. B.O.C. 1997 117(4) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 117 No. 4 Published 12 December 1997 


The eight hundred and sixty-ninth meeting of the Club was held on Tuesday 17 June 
1997, at 6.30 p.m. 21 Members and 9 guests attended. 

Members present were: The Rev. T. W. GLapwiIn (Chairman), R. W. Woops 
(Speaker), Miss H. Baker, Captain Sir Thomas BarLow Bt. RN, J. W. BarrINGTON, 
Captain M. K. Barritt RN, P. J. BELMan, D. R. CALDER, Cdr M. B.-CasEMENT RN, 
R. B. CurLpress, S. J. FARNSworTH, D. J. FisHER, D. GriFFIN, J. A. JOBLING, Dr 
J. B. Kerstey, D. J. Montier, Mrs A. M. Moore, R. G. Morcan, Dr W. G. Porteous, 
N. H. F. STONE and Mrs F. E. Warr. 

Guests attending were: Mr M. Apams, Mrs A. Brown, Mrs C. R. CasEMENT, Mrs S. 
CHILDREss, Mrs J. M. Giapwrn, Mrs S. GrirFin, Mrs S. Kerstey, Mrs M. MONTIER 
and Mr J. Warr. 

After dinner, Mr Robin Woods spoke on the Atlas of Breeding Birds of the Falkland 
Islands, illustrating his talk with slides of birds, habitats and data from the Atlas. 
Together with his wife Anne, who jointly edited the book, he lived in Stanley for six 
years to 1963, while he worked for the British Antarctic Meteorological Service. His first 
book on Falkland birds was published in 1975. The Falklands consist of two main 
islands, and about 780 smaller islands. They cover 4,700 sq. miles, extend 150 miles, 
east to west, but the population is only about 2,200 of whom two-thirds live in the 
only town of Stanley. A further 2,000 military personnel are based at Mount Pleasant 
Airport. 

There are no native trees but Cupressus macrocarpa and shrubs have been planted in 
Stanley and at most of the 80 or more remote farm settlements, providing good shlter for 
gardens and birds. The native Tussac grass Parodiochloa flabellata formerly grew up to 
3 m (10 feet) tall in dense coastal fringes and was mistaken for palm trees by early visitors. 
Uncontrolled grazing by cattle and sheep has destroyed over 80% in total, and almost all 
Tussac on East and West Falkland. It survives as a single plant community on ungrazed 
offshore islands where it supports about 30 bird species. A century ago, some farmers saw 
its value as fodder or shelter and replanted coastal paddocks. Bird density inland, away 
from ponds or streams, is very low. Most of the country is open moorland with rock 
outcrops. Roads of crushed stone have been constructed recently and the Government 
Air Service provides transport to about 30 inhabited settlements with airstrips, but travel 
to the outer islands is difficult. Kidney Island, which has thousands of colonial seabirds, 
and was declared as the first Falkland Island Government Nature Reserve in 1964, can 
only be reached by sea. 

Returning in late 1983 to study passerines in Tussac grass, Robin collected records of 
breeding birds from residents. Ideas for an Atlas were discussed with members of the 
Falkland Islands Trust in Stanley, and the Falkland Islands Foundation in UK. Both 
organisations, which later merged to become Falklands Conservation, supported the 
project, and record forms reached the Falklands for the 1984/85 season. Robin described 
the development of the project over ten breeding seasons, with examples of local 
publicity, data collection and analysis. Records were received from residents, military 
personnel in the Islands, observers on Naval patrol ships, and tourists. One visiting 
scientist was funded to cover the more inaccessible parts of the main islands. 
Observations varied according to the detectability of species, observer skills and 
topography. Penguins were highly visible while elusive nocturnal petrels were all 
under-recorded. In the Atlas, survey results and notes on habitat and breeding are 
compared with historical data. The distribution of most species is related to the main 
topographical features such as the number of ponds, the amount of coast and the attitude 
of land within each 10 km grid square. 

Some interesting species were discussed. These included the Upland Goose 
Chloephaga picta which occurs in more squares than any other species. The closely 
related Ruddy-headed Goose Ch. rubidiceps is also widespread, but has recently been 
classified as Near-threatened by BirdLife International because the continental 
population has declined drastically. The endemic Falkland Flightless Steamer Duck 


Meetings 242 Bull. B.O.C. 1997 117(4) 


Tachyeres brachydactyla and the Kelp Goose Ch. hybrida are both thriving in coastal 
squares. Three species of gulls show very different levels of abundance. 

The Barn Owl Tyto alba was proved to breed for the first time during the survey. It 
nests in dense European Gorse Ulex europeus which was introduced about a century ago 
as cattle fencing. Pellet analysis shows that the introduced feral House Mouse Mus 
domesticus is the main prey animal. In contrast, a very tame endemic race of the Blackish 
Cinclodes Cinclodes antarcticus (the Tussacbird), cannot survive with rats, mice or cats 
but is numerous on coasts of outer islands. Similarly, the endemic Cobb’s Wren 
Troglodytes cobbi, first described at a B.O.C. meeting in 1909, is found only on offshore 
Tussac islands without mammalian predators, and is now classed as Vulnerable. Its 
susceptibility is probably linked to its habit of feeding and nesting at or below ground 
level. Records of feral cats Felis catus, Common Rats Rattus norvegicus and House Mice 
show the wide distribution of these introduced predators. 

The Falklands may hold the majority of the world population of the Black-throated 
Finch Melanodera melanodera. It is present in good numbers, though classed as 
Near-threatened because continental birds are declining, through destruction of 
habitat. A very unusual, tame and inquisitive bird of prey, the Striated Caracara 
Phalcobaenus australis, also Near-threatened, is recovering slightly after more than a 
century of persecution as a pest of sheep farming. It breeds on offshore islands with 
albatross and penguin colonies, and has a very restricted distribution in extreme southern 
South America. The Falklands, with about 600 pairs, hold the majority of its population. 

The Atlas discusses the conservation implications of environmental factors, including 
offshore fishing, probable oil extraction developments and imminent revision of local 
wildlife protection legislation, for all breeding species. A recent collection of bird bones 
from peat deposits on West Point Island may give evidence of the occurrence of species 
now absent, and radiocarbon dating would make an impressive difference to knowledge of 
the history of Falkland Islands birds. 

In thanking Robin for his talk, the Chairman introduced Mrs Anne Brown, Secretary 
of Falklands Conservation (Robin’s guest), who joined in the subsequent lively question 
and discussion period. 


The eight hundred and seventieth meeting of the Club was held on Tuesday 15 July 1997 
at 6.15 p.m. 27 Members and 9 guests attended. 

Members present were: The Rev. T. W. GLADWIN (Chairman), Miss H. Baker, C. R. 
BarLow, J. W. BarrINGcTON, D. M. BrapD.ey, P. J. BULL, Cdr M. B. CASEMENT RN, 
Dr R. A. CHEKE, Dr R. A. F. Cox, R. B. CHILDREss, D. J. FISHER, F. M. GAUNTLETT, 
A Gisps, D. GRIFFIN, J: A. JOBLING, R. H. Kett.e, M. B. Lancaster, D. J. MONTIER, 
Mrs A. M. Moore, Mrs M. N. MutLtier, Dr R. P. Prys-Jongs, N. J. REDMAN, R. E. 
Scott, N. H. F. STONE and C. W. R. STorREy. 

Guests attending were: T. APPLETON (Speaker), M. BrapLey, Mrs J. Butt, Mrs S. 
CuILprRess, Mrs J. M. GLapwin, Mrs M. H. Gaunt ett, P. J. Moore, C. A. MULLER 
and Mrs S. STONE. 

On completion, Tim Appleton gave an illustrated talk entitled “‘From Greenfields to 
Ramsar’’ about the history and development of Rutland Water. 


The eight hundred and seventy-first meeting of the Club was held on Tuesday 16 
September 1997 at 6.15 p.m. 25 Members and 13 guests attended. 

Members present were: Mrs A. M. Moore (Chairman), G. E. GREEN (Speaker), Miss 
H. Baker, J. W. BarriInNcTON, P. J. BELMaNn, I. R. Bishop, D. R. CALDER, Cdr M. B. 
CASEMENT RN, F. M. GauntTLett, D. GriFFIn, J. A. JoBLinc, R. H. KetTt ie, Dr C. F. 
Mann, D. J. Montier, Mrs M. N. Mutter, R. G. Morean, P. J. OLtver, Dr W. G. 
Porteous, Dr R. P. Prys-Jonsgs, N. J. Repman, P. G. W. Sataman, Dr D. W. Snow, 
S. A. H. Statuam, N. H. F. STONE and G. THomas. 

Guests attending were: Ms G. Bonuam, Mrs J. B. CaLprer, D. GaNpy, T. Evans, Mrs 
M. H. GauntietTtT, Mr & Mrs D. B. ILEs, Mrs M. Montisr, P. J. Moore, R. RAnrtT, 
Dr B. M. Rocers, Mrs K. SaLaman, and Mrs B. K. Snow. 

On completion, Graeme Green gave an illustrated talk entitled “‘Cotingas and their 
niche in the neotropical avifauna’’. 

The cotingas as a group reflect the diversity which is typical of the Neotropical 
avifauna in general. The geological history of the Americas is such that there are many 


Meetings 243 Bull. B.O.C. 1997 117(4) 


centres of avian endemism, both lowland and montane, and many cotingas are endemic to 
just one of these areas. This endemicity and resultant dependence on one discrete area 
makes these cotingas vulnerable to habitat modification. This factor has perhaps reached 
its nadir in the rampant destruction of the Atlantic forests of Brazil, Argentina and 
Paraguay, where perhaps only 4% of primary forests remain. Whether this is sufficient 
habitat for viable populations to survive may be answered sooner rather than later. Also, 
it is not just these discrete areas of endemism where habitat modification is such a 
problem; the whole of the eastern slope of the Andes is under threat of complete forest 
clearance outside protected areas, particularly within the elevational range suitable for 
agriculture based on cash crops such as sun coffee and marijuana. 

Hindering our understanding of the possible consequences to the Neotropical avifauna 
in general, and cotingas in particular, of this habitat modification is the fundamental lack 
of baseline data for many Neotropical birds. The ground-breaking efforts of many 
ornithologists such as Frank Chapman, Alexander Skutch, and others, has developed into 
a modern field-orientated ornithological movement. This growing cadre of field 
ornithologists may become crucial in gathering sufficient information on the birds of the 
Neotropics to help decision-makers to avert further extinctions. 

The increasing understanding of the crucial role that vocalisations play in the lives of 
Neotropical birds has led to vast collections of taped vocalisations being housed in 
various institutions, where they are accessible to researchers. There are also more 
commercial tapes appearing on the market, and these can also be extremely useful in 
increasing researchers’ knowledge of the avifauna. Cotingas exhibit a wide repertoire of 
vocalisations, ranging from the woeful and near-mute to the spectacular. Knowledge of 
their vocalisations is a useful tool in mapping distributions of these birds, as many are 
attracted to tape playback, or whistled imitations, of their vocalisations. For example, the 
Elegant Mourner Laniisoma elegans (the erstwhile Shrike-like Cotinga), has a distribution 
centred on the Atlantic Forests of Brazil from where, it is speculated, it colonised the 
eastern slope of the Andes of Peru, Ecuador, Colombia and Venezuela (Snow 1982). The 
vocalisation of the Atlantic Forest taxon of this species, L. e. elegans is now widely known 
(indeed, it is now commercially available), and this has led to more data being gathered 
about the taxon, which can be extremely difficult to observe, during fieldwork in the 
region. Consequently, the belief among many ornithologists is that this taxon is both 
more widespread, and less rare, than formerly considered. 

The Atlantic Forests of Brazil harbour several other cotingas with interesting 
vocalisations. Anyone who has heard the ululating, high-pitched and far-carrying whistle 
of the Black-and-Gold Cotinga Tyuca atra will not forget its ventriloquial quality. 
Another characteristic (and beautiful) cotinga of the region, the Hooded Berryeater 
Carpornis cucullatus has a vocalisation which has earned it the onomatopoeic name 
“coracacho’; indeed it is perhaps the characteristic call of the wetter forests of the Serra do 
Mar. The rediscovery in the region in 1996 of the feared-extinct Kinglet Calyptura 
Calyptura cristat is a fitting reward for the efforts of the fieldworkers in the region, and 
epitomises the requirements for solid, and at times monotonous data-gathering in the 
cause of conservation of these wonderful and spectacular birds. 


Erratum: 

In the account by Prof. Richard Chandler Bulletin 117 (3), p. 158, a sentence was 
omitted. The following sentence should be inserted before the last sentence in paragraph 
2: “The two forms also differ in both orbital ring (narrow and yellow-orange in 
fuliginosus, fleshy, broad and orange-yellow in ophthalmicus) and claw colour (orange in 
fuliginosus and black in ophthalmicus).” 


P. A. Clancey 244 Bull. B.O.C. 1997 117(4) 


Subspeciation in Layard’s Tit-babbler of the 
southwestern Afrotropics 


by P. A. Clancey 
Received 18 May 1996 


The tit-babblers Parisoma subcaeruleum and P. layardi form a small 
unit of sympatric species endemic to the South African Sub-Region, 
where they affect the Acacia savanna biome. They are thought to be 
closely related to the Old World warblers of the genus Sylvia. While 
both South African species are broadly sympatric, P. layardi is the less 
common of the pair and is confined to the extreme west and the south 
of their joint range, which extends from southwestern Angola, the 
mid-Zambezi drainage and the plateau of Zimbabwe, south to the Cape 
and western Natal in the Republic of South Africa. The other species 
currently treated as congeneric are P. buryi of southwestern Arabia, and 
P. lugens and P. boehmi of the northeast and eastern parts of Africa. P. 
buryt is monotypic, while P. lugens and P. boehmi are moderately 
polytypic, as are subcaeruleum and layardi. While the two southern 
African forms are sympatric and share the same Acacia biome, they are, 
nevertheless, moderately differentiated ecologically, with layardi found 
in broken, hill country with tracts of rocky terrain. 

In the west of its range, /ayardi is present along the western seaboard 
at sea level, but in the extreme east, in the highlands of Lesotho, it 
affects high country, breeding to elevations in excess of 2500 m and 
reaching the alpine summit of the high Drakensberg. From the 
ecological point of view it is significant that in the west of the range 
precipitation levels are low, being c. 120 mm annually, whereas in the 
extreme east the birds espouse an environment which experiences a 
rainfall of c. 600-1500 m. 

Traylor, in his 1986 appraisal of subspeciation in P. subcaeruleum and 
P. layardi, admitted four subspecies for both species. While I have no 
cause for cavil over the four subspecies admitted in subcaeruleum, the 
recognition of four in P. layardi calls for reconsideration, and is 
critically reassessed hereunder. 

With its limited range, P. layardi has invited a limited measure of 
interest on the part of systematists. The first worker to describe a 
form in P. layardi, which was named initially from Clanwilliam in 
1862 by Hartlaub, was Vincent (1948) who, on the basis of a limited 
material of eight specimens from the Lesotho highlands and the small 
Transvaal Museum series of ten then available, separated the Lesotho 
birds on the basis of colour and bill-length characters. Later, 
Winterbottom (1958) separated a race on a comparable range of 
colour variables, based on five specimens from the coastlands of the 
western Cape and nineteen from a range of localities to the northeast 
in the lower Orange R. basin of the Richtersveld. Both descriptions 
compared a dark blue-grey-backed form with a lighter, more 
olivaceous-backed form (the nominate), no attempt being apparently 


P. A. Clancey 245 Bull. B.O.C. 1997 117(4) 


made to bring the two dark grey variants together in the comparisons. 
The geographical variation in the present Parisoma is relatively 
simple, grouping readily into two classes, one dark bluish-grey with 
white in the wings and over the lores, and the other olivaceous grey, 
without white in the wings and over the face. Birds in the former 
group also frequently exhibit a prominent white supra-loral spot. 
Strangely, neither Vincent nor Winterbottom alluded to the strong 
development of white over the wings and the face. This was 
particularly strange in the case of Vincent’s highland material from 
Lesotho which he described as P. /. barnes, these all strongly marked 
with white in the wings, but in the western Cape where comparable 
birds are seemingly confined to the dune country, of which 
Winterbottom had but five specimens, birds of both forms come into 
very close proximity to one another. 

The finding of two widely sundered populations showing closely 
comparable suites of characters in P. layardi follows a comparable 
trend in a number of Cape endemics, notably Promerops cafer/ 
P. gurneyi, Chaetops frenatus/Ch. aurantius and Pseudochloroptila 
pseudochloroptila/P. symonst. However, unlike the listed endemics, the 
forms here discussed are not even subspecifically differentiated. In the 
present case there is no ready ecological interpretation, as western birds 
occupy scrub along the southwestern Cape coast and eastern birds are 
typical for the species. 

The second variant population, differentiated from the dark bluish birds 
already dealt with, is more olive-tinged over the dorsal surface and wings, 
and moreover lacks white over the remiges and usually the face, which 
surfaces are olive-grey. On the underside they differ little from the blue- 
grey birds though tending to show more white medio-ventrally. To such 
elements Winterbottom applied the name P. /. aridicola, the type a bird 
taken at Noisabis in the Richtersveld. A critical examination reveals that 
birds agreeing taxonomically with aridicola extend far to the south of the 
Richtersveld and the basin of the lower Orange R., where they lie in 
juxtaposition with dark birds confined to the dunes along the coast to the 
west. Birds agreeing in such characters were collected at a range of localities 
extending from Kamiesberg to Calvinia and Citrusdal. Some were identi- 
fied by Winterbottom in 1958 as aridicola, which raises the question of the 
status of aridicola as being other than a junior synonym of layardi, proposed 
on a Clanwilliam specimen. In essence, this would deprive the present 
layardi of a name, which can however be met by seeing the latter as 
congruent with the eastern highland taxon as part of P. 1. barnesz. 

A further name requires to be considered, P. 1. subsolanum Clancey, 
1963: Aprilskraal siding, near Molteno, northeastern Cape. This is not 
a particularly sharply defined subspecies, but is seen as a connecting 
link between the two dark well-characterised southern populations of 
P. l. barnesi in having the breast and sides darker, yet lacking any 
marked white in the wings and face, and can be gainfully employed for 
the population of “‘nominate layardi-type’’ birds present throughout 
the southern mountains and Karoo of the Cape. 

Nominate P. /. layardi extends from the central and southern parts of 
Namibia south to the northwest of the Cape in the basin of the lower 


P. A. Clancey 246 Bull. B.O.C. 1997 117(4) 


Orange R., south to about Port Nolloth on the coast and in the south in 
the interior of Little Namaqualand to Clanwilliam, Calvinia and 
Citrusdal, thence northeast to the middle Orange R. Both nominate 
layardi and subsolanum are separable in the entire lack of white in the 
wings and face, such parts being olivaceous grey (about the Deep 
Mouse Gray of Ridgway). 

‘Three subspecies can be admitted in the present species: P. 1. barnesi, 
P.1.layardi and P. 1. subsolanum, the first-named in two widely sundered 
populations. They are based entirely on plumage characters, no signifi- 
cant differences having been detected in their linear measurements. 


Parisoma layardi barnesi Vincent 1948, Bull. Brit. Orn. Cl. 68, p. 145: 
Lekhalabaletsi R. valley above confluence of the Jareteng and 
Lekhalabaletsi Rivers, Lesotho at 29°17'S, 29°24’E, 8900 ft. 

Upper-parts and wings dark bluish leaden grey (about Dark Neutral 
Gray of Ridgway), and occasionally with a well-defined narrow white 
superciliary stripe over the lore. Below, white with narrow blackish 
streaking over the fore-throat and with the breast and sides moderately 
washed with light neutral grey. ‘The wings with the outer vanes of the 
remiges broadly edged with white, this carried to the tertials, forming a 
well-defined white wing-stripe. 

Range. Occurs in two widely sundered populations, the eastern one 
confined to the highlands of Lesotho and adjacent northeastern parts of 
Cape Province (Drakensberg Range and the interior ranges of the 
Maluti Mts). A second undifferentiated population is present along 
the western Cape where it occurs in maritime vegetation from the 
Port Nolloth district in the north, south narrowly to the southwestern 
Cape. Localities:- W. Cape: Port Nolloth, Wallekraal, —TTwee Dam 
(Springbok), Karoopoort (Koue Bokkeveld), Kliprand 27 m N. of Garies, 
Bitterfontein, Lootsberg Pass, Ysterfontein, Melkbos, Muizenberg. E. 
Cape: Naude’s Nek Pass, Strandfontein. Lesotho: Marakabeis, mountains 
E. of Maseru. . 

Remarks. The type-specimens of P. layardi (Clanwilliam) and P. 1. 
aridicola were made available for comparison by the South African 
Museum, Cape Town. The layardi type had been mounted and its 
coloration is now wholly unreliable. 

Parisoma layardi layardi Hartlaub 1862, Ibis (1)4, p. 147: Zwartland, 
Malmesbury district; corrected to Clanwilliam, western Cape, by 
Winterbottom 1957, Ostrich 28, p. 235. 


Parisoma layardi aridicola Winterbottom 1958, Bull. Brit. Orn. Cl. 78, 
p. 148: Noisabis, Richtersveld, northwestern Cape. 

Upper-parts and wings distinctly lighter, less dark bluish-grey, than 
in P. 1. barnesi, being about Deep Mouse Gray of Ridgway, ard without 
white over the wings and face, the outer vanes of the flight-feathers 
greyish olive-buff. 

Compared to subsolanum differs in the starkly whiter ground to the 
fore-throat, and the greater extent of the mid-ventral white. Upper- 
parts lighter olivaceous-grey. | 

Range. The highlands of Damaraland, Namibia, south in the west of 
Namibia, inland of the Namib Desert, to the Richtersveld and basin 


P. A. Clancey | 247 Bull. B.O.C. 1997 117(4) 


of the lower Orange to the coast at Port Nolloth. In the interior and east 
of Little Namaqualand, south to Clanwilliam and the Citrusdal areas, 
and east to Upington, Carnarvon and Victoria West. Localities:- 
Namibia: Great Karas Mtns. Cape: Goodhouse, Namees, Noisabis, 
Pofadder, Upington, Port Nolloth, Carnarvon, Kamiesberg, Garies, 
Vanrhynsdorp, Lokenburg (Calvinia), Clanwilliam, Citrusdal, and 
localities immediately to the east of southern N. Cape. 

Remarks. One or two specimens from near Garies and Vanrhynsdorp 
in Little Namaqualand show a small amount of white in the remiges, 
revealing a measure of intergradation towards P. l. barnesi, although 
retaining the diagnostic lighter mouse-grey to the upper-parts and 
wings. 


Parisoma layardi subsolanum Clancey 1963, Durban Mus. Novit. 6, 
p. 253: Aprilskraal siding, Molteno, northeastern Cape. 

Compared with P. layardi differs in being less starkly white over the 
ground of the fore-throat, and with the breast darker, more buff, less 
whitish, and with the sides of the body and the flanks much darker 
grey, with less white medio-ventrally. Upper-parts and wings slightly 
darker, especially over the pileum, but not as dark and blue as in P. J. 
barnes. 

Range. The highlands of the southwestern Cape, extending east 
through the southern mountain ranges and adjacent regions of the 
Karoo to the eastern and northern Cape and the Orange Free State. 
Localities:- Hanover, Williston, Laingsburg, Oudtshoorn, Campher- 
spoort (East London), WHuisrivierpas, Danebury, Graaff-Reinet, 
Rossouw, Murraysburg, Jamestown, T’eebus, Lelykpoortjie, Molteno, 
Sterkstroom, Griquatown and Aliwal North. 

Remarks. This subspecies lies close to the. nominate one in lacking a 
starkly white wing-stripe and loral mark, but its darker ventral parts in 
a sense link the two dark forms of P. |. barnesi lying to the east and west 
of it. 


Acknowledgements 


For the loan of additional material I am indebted to the Ornithologist of the East London 
Museum (Mr C. J. Vernon) and the Director of the South African Museum, Cape Town. 
Just on ninety specimens were available for the present research, including three Types. 


References: 

Clancey, P. A. 1963. Notes, mainly systematic, on some birds from the Cape Province. 
Durban Mus. Novit. 6: 252-254. 

Ridgway, R. 1912. Color Standards and Color Nomenclature. The author, Washington. 

Traylor, M. A. 1986. Check-list of Birds of the World. Vol. 11. Museum of Comparative 
Zoology, Harvard. 

Vincent, J. 1948. New races of a Tit-babbler and a Lark from the Basutoland Mountains. 
Bull. Brit. Orn. Cl. 68: 145. 

Winterbottom, J. M. 1958. A new subspecies of Parisoma layardi Hartlaub. Bull. Brit. 
Orn. Cl. 78: 148-149. 


Address: Dr P. A. Clancey, Durban Natural Science Museum, P. O. Box 4085, Durban 
4000, South Africa. 


© British Ornithologists’ Club 1997 


N. Krabbe et al. 248 Bull. B.O.C. 1997 117(4) 


Range extensions of cloud forest birds from 
the high Andes of Ecuador: new sites for rare 
or little-recorded species 


by Niels Krabbe, Bent Otto Poulsen, Amy Frolander & 
Orfa Rodriguez Barahona 


Received 5 August 1996 


A comparative survey was made of the avifauna at ten sites with 
humid forest in the temperate zone of Ecuador (Fig. 1) in January 
and February 1995 and 1996. The major comparison of diversities 
will be published elsewhere, but we here give the records that 
represent altitudinal or southward range extensions, and, for rare or 
little-recorded species, we also list the sites where we found them. 
Most records are from within the study sites, which all ranged from 
3000 to 3350 m in altitude, but some are from above or just below the 
sites, and for some records, additional observations have been 
included. 

Abbreviations of localities are (province and locality coordinates in 
parenthesis): East slope (north to south): Oyacachi=below Oyacachi 
(Napo: 00°13'S, 78°02’W); Anatenorio=Rio Anatenorio (Napo: 
00°59’'S, 78°17'W); Matanga=Paramos de Matanga (Morona- 
Santiago: 03°16’S, 78°54'W); ‘'Toledo=Cerro ‘Toledo (Zamora- 
Chinchipe: 04°23’'S, 79°06’W); Lagunillas=Cordillera Las Lagunillas 
(Zamora-Chinchipe: 04°46'S, 79°25'W). West slope (north to south): 
Intag=Intag (Imbabura: 00°20’N, 78°25'W); Corazon=Volcan 
Corazon (Pichincha: 00°33’'S, 78°43’W); Salinas=10 km northwest of 
Salinas (Bolivar: 01°21’S, 79°05’W); Chaucha=above Chaucha 
(Azuay: 02°52'S, 79°23'W); Mazan=Rio Mazan (Azuay: 02°52'S, 
79°07'W). 

Rio Mazan is here considered west slope (it lacks 22 species restricted 
to the east slope), but its avifauna includes some eastern and 
southwestern (as well as endemic) elements. 

Other abbreviations used in the text: ANSP (Academy of Natural 
Sciences of Philadelphia), MECN (Museo Ecuatoriano de Ciencias 
Naturales, Quito), ZMUC (Zoological Museum, University of 
Copenhagen), NK (Niels Krabbe) and BOP (Bent Otto Poulsen). 


Altitudinal records 
HIGHLAND TINAMOU Nothocercus bonapartei 


One was observed at close range for several minutes on two 
occasions at 3075 m at Oyacachi. Previously only known up to c. 
2200 m in Ecuador and Colombia (Robbins et al. 1987, Hilty & 
Brown 1986), but recorded to 2500m in Venezuela (Meyer de 
Schauensee & Phelps 1978). 


N. Krabbe et al. 249 Bull. B.O.C. 1997 117(4) 


ie = o Oyacachi 


Corazon ~ a : oi 
Mion’). 18 


na tpn i é Anatenorio 


: =~ as: tf : 
Salinas @ See 


San, 


Ta ad fae tal 
Riobamba = i 
Ro? 


» 


Guayaquil 


Se Al 
So 
= 
- 
a? cow 
m— aS t 3 
~_ ow 
Se 


a wate 


ies 


Chaucha: 70° “*,Cuenca 


AmaluZa ss 


ae / 
Ne Laginitlas 


Figure 1. Map of Ecuador showing the five eastern slope (rhombuses) and the five 
western slope sites (large circles) investigated in this study. The two largest cities 
(squares) and some principal Andean cities (small circles) are also shown. The 3000 m 
contour is stippled. 


ROADSIDE HAWK Buteo magnirostris 


A pair was seen at Oyacachi and a pair at Anatenorio, both at 3000 m. 
Previously known up to c. 2500 m (own obs.). 


AMETHYST-THROATED SUNANGEL Heliangelus amethysticollis 

A single male was seen at 3025 m at Lagunillas. Previously known up 
to 2400m in Ecuador (also at Lagunillas) (M. B. Robbins pers. 
comm.), but recorded up to 2600m in immediately adjacent Peru 
(Parker et al. 1985), to 3000 m in Colombia and Venezuela (Hilty & 
Brown 1986, Meyer de Schauensee & Phelps 1978), and sighted by NK 


N. Krabbe et al. 250 Bull. B.O.C. 1997 117(4) 


to 3200m in Cuzco, Peru (Fjeldsa & Krabbe 1990). Its lower 
altitudinal limits throughout lie at 1800-2000 m. 


EMERALD TOUCANET Aulacorhynchus prasinus 

To 3250m below Oyacachi, whence come three old specimens 
without exact altitudes given (Chapman 1926). Previously only 
definitely known up to 2750 m in Ecuador (Krabbe 1991), to 2200 m in 
Peru (Parker et al. 1985, Fjeldsa & Krabbe 1990) but recorded to 
3000 m (once to 3700 m) in Colombia (Hilty & Brown 1986). 


BARRED ANTTHRUSH Chamaeza mollissima 
A pair was observed at 3075 m at Oyacachi. Previously not known 


above 2300 m in Ecuador (own obs.), but in Colombia known to 
3100 m (Hilty & Brown 1986). 


BARRED BECARD Pachyramphus versicolor 

Recorded to 3000 m at Oyacachi and to 3050 m on Toledo. The only 
previous record from above 2850 m in Ecuador is of a bird mist-netted 
and photographed (BOP, T. Lessge) at 3500 m on Volcan Pichincha 
(Fjeldsa & Krabbe 1990). In Colombia not reported above 2600 m 
(Hilty & Brown 1986), but known up to 2950 m in northern Peru 
(Parker et al. 1985). 


SLATY-BACKED CHAT-TYRANT Ochthoeca cinnamomeiventris 

At Lagunillas singing birds were observed to 3320 m. Previously 
known up to 3000m (Fjeldsa & Krabbe 1990). Occasionally strays 
higher: in Cordillera de Los Llanganates, southern Napo Province, one 
bird appeared at 3600 m in early May 1992, sang intensely for a few 
hours, and then flew down slope, not to be seen or heard there again 
over the next days (NK). 


GREATER PEWEE Contopus fumigatus 


To 3000 m at Oyacachi. Previously only known up to c. 2500 m in 
Ecuador (Krabbe 1991), but locally to 3000 m in Colombia (Hilty & 
Brown 1986). 


BLACK-COLLARED JAY Cyanolyca armillata 


Found to 3150 m at Oyacachi, whence originates a specimen taken in 
1950 (ANSP) without an exact altitude. Otherwise only known in 
Ecuador from two specimens taken at Rio Pun in northeastern Carchi 
or adjacent Sucumbios Province (Salvadori & Festa 1899) without 
altitudes given, but probably between 2200 and 2600m, and a 
specimen taken by J. C. Matheus at 2320 m at La Alegria below Pun 
(MECN) and sightings down to 2050 m below La Alegria (NK). The 
two species (C. armillata and Turquoise Jay C. turcosa) may thus 
replace each other altitudinally in the Rio Pun valley, where the east 
slope forest meets the interandean forest. On the interandean slope not 
far south of the Rio Pun valley C. turcosa occurs up to 3350 m (Robbins 
et al. 1987). At Oyacachi, however, we found C. armillata up. to 
3150 m, and no C. turcosa, so they apparently exclude each other 


N. Krabbe et al. 251 Bull. B.O.C. 1997 117(4) 


locally. In the next valley to the south of Oyacachi (Papallacta), C. 
turcosa occurs on the entire slope, at least down to below 2500 m (own 
obs.). Hilty & Brown (1986) reported C. turcosa from both slopes of 
Narino, Colombia at 2600-3000 m, but did not provide any details of 
altitudinal distribution in eastern Narino, where the two species 
overlap. Vocally, they differ distinctly (own obs.). 


GLOSSY-BLACK THRUSH Turdus serranus 

Mostly known from below 2800 m. Singing up to 3150 m at Intag 
and Anatenorio, and to 3350m at Oyacachi. In mid-March 1996 
several singing birds were tape-recorded as high as 3750 m at Loma 
Yanayacu on Volcan Pichincha (NK), where, despite many visits by 
birdwatchers, they had not been recorded before. 


RUFOUS-BROWED PEPPERSHRIKE Cyclarhis gujanensis 
One singing bird of the southeastern subspecies contrerasi was 
observed and tape-recorded at 3100 m on Paramos de Matanga. This 


form had previously not been found above 1900 m in Ecuador (own 
obs.). 


BROWN-CAPPED VIREO Vireo leucophrys 


Two birds, clearly a pair, were observed and their song and calls 
tape-recorded at 3150m at Oyacachi. Previously not known above 
2650 m in Ecuador (Krabbe 1991). In Colombia recorded up to 2800 m 
(Hilty & Brown 1986). 


GRASS-GREEN TANAGER Chlorornis riefferit 

Observed up to 3175m at Intag and to 3500m on Corazon. 
Previously known up to 2900m in Ecuador (Krabbe 1991). In 
Colombia rarely recorded to 3300 m (Hilty & Brown 1986). 


CHESTNUT-CAPPED BRUSH-FINCH Buarremon brunneinucha 

Seen up to 3150m on Toledo. The only previous record above 
2750 m in Ecuador is of a bird that died in a mist-net at 3150 m at 
Acanama, northern Loja Province (R. Williams, J. Tobias; skeleton in 


Salango Museum, NK preparation). No Buarremon torquatus was 
recorded at Toledo (see Remsen & Graves 1995). 


Southward range extensions 
RUFOUS-BANDED OWL Ciccaba albitarsus 


Found at all five sites on the west slope, where it had previously not 
been found south of Pichincha (Chapman 1926). 


RUFOUS-BELLIED NIGHTHAWK Luyocalis rufiventris 


Found at Chaucha. Previously known only south to western 
Chimborazo on the west slope (Berlepsch & Taczanowski 1884). There 
is an unpublished record (by NK) from near Molleturo just north of 
Chaucha. 


N. Krabbe et al. 252 Bull. B.O.C. 1997 117(4) 


BUFF-WINGED STARFRONTLET Coeligena lutetiae 


Found at Salinas. Only previous record from south of Pichincha on 
the Pacific slope is from western Cotopaxi (Krabbe 1991). 


ANDEAN TIT-SPINETAIL Leptasthenura andicola 

Found at 3250-3300 m at Lagunillas. It had been overlooked at this 
locality during earlier surveys (ANSP team pers. comm., own obs.), 
and undoubtedly also at Cerro Chinguela in immediately adjacent 
Peru (Parker et al. 1985). In Ecuador the species had previously 
been recorded south to Cajas in Azuay (Gretton in Robinson 1987, 
own obs.) (subspecies andicola), some 200km north of Lagunillas. 
The northernmost Peruvian record (subspecies peruviana) is from 
Cordillera Blanca, Ancash, some 400 km south of Lagunillas (Fjeldsa & 
Krabbe 1990). The Lagunillas birds may thus represent an undescribed 
subspecies. 


BARRED FRUITEATER Pipreola arcuata 


Recorded at Salinas. The southernmost previous record on the west 
slope is from Pichincha (Chapman 1926). 


SLATY-BACKED CHAT-TYRANT Ochthoeca cinnamomeiventris 

In Cordillera de Chilla, northwesternmost Loja Province, we 
observed this species between Selva Alegre and Manu. We also found it 
to 3050 m at Rio Mazan. On the west slope previously not reported 
from south of Chimborazo (Chapman 1926). It has undoubtedly been 
overlooked on the Pacific slope in Azuay. 


BLACK-CAPPED TYRANNULET Phyllomyias nigrocapillus 


Found at Salinas and Chaucha. Previously not recorded south of 
Pichincha on the west slope (Chapman 1926). 


RUFOUS WREN Cinnycerthia unirufa 


Recorded at Salinas. On the west slope, previously known south to 
western Cotopaxi (Krabbe 1991). 


BLACK-BACKED BUSH-TANAGER Urothraupis stoltzmanni 

A flock of ten was observed in treeline scrub down to 3300 m at 
Matanga. Previously unrecorded south of Zapote-Najda mts. (Fjeldsa 
& Krabbe 1986). Like Crescent-faced Antpitta Grallaricula linetfrons 
(see below), it undoubtedly ranges along the entire eastern Andes of 
Ecuador south to Rio Zamora. 


New localities for rare or little-recorded species 
IMPERIAL SNIPE Gallinago imperialis 


Roding at all five eastern, and the two northernmost western sites. At 
five localities it had not been recorded earlier. In Ecuador the species 
appears to be continuously distributed along the entire east slope and, 
though as yet unrecorded in western Carchi, along the west slope south 
to Volcan Iliniza. The twelve Ecuadorian localities known are: 


N. Krabbe et al. 253 Bull. B.O.C. 1997 117(4) 


1. 
2. 


10. 


11. 
£2) 


Intag, Imbabura, 00°20'N, 78°25’W, 3330-3350 m: present study 
(5-11 January 1995). 

Volcan Pichincha, Pichincha, 00°08’S, 78°35’W, 3600-3800 m: 
Krabbe (1991, 1992). Roding only in the rainy season (October to 
May). NK collected a male specimen here on 8 March 1992 
(MECN 6015). Although roding, it had only medium-enlarged 
testes. It had worn wings and tail, and was in heavy body moult. 
Measurements in mm: wing chord 155, wing flat 160, tail 55, bill 
from feathers 76, tarsus 35. Body mass 196g. Only three other 
specimens exist, two of them without data (Terborgh & Weske 
1972). 

Volcan Corazén, Pichincha, 00°33’S, 78°43’W, 3150-3750 m: 
present study (17—24 January 1995). 

Cerro Mongus, Carchi, 00°27'N, 77°52’W, 3200-3500 m: L. 
Petersson tape-recordings 5—6 July 1995 (Petersson 1996). 

Below Oyacachi, Napo, 00°13’S, 78°02’W, 3350 m: present study 
(10-14 January 1996). 

Rio Anatenorio, Napo, 00°59’S, 78°17'W, 3300-3350 m: present 
study (18-22 January 1996). 

Cordillera de los Llanganates, Napo, 01°06’S, 78°18’W, 3500 m: 
NK tape-recordings (26-28 May 1992). 

Paramos de Matanga, Morona-Santiago, 03°16’S, 78°54’W, 
3100-3300 m: NK _ tape-recordings (14-15 November 1992); 
present study (6-10 February 1996). 

Acanama, Loja, 03°40’'S, 79°14’°W, 3200 m: NK tape-recording 
(6 November 1992). 

Cajanuma, Loja, 04°06’S, 79°09’W, 2800-3000 m: Pearman (1994) 
(7-10 December 1990); tape-recordings by R. Williams (January— 
February 1991); individual seen foraging and photographed by 
Poulsen (1993) (March 1992); chick found by M. Lysinger 
(February 1993). 

Cerro Toledo, Zamora-Chinchipe, 04°23'S, 79°06’'W, 3030- 
3350 m: present study (20-24 February 1996). 

Cordillera Las Lagunillas, Zamora-Chinchipe, 04°46'S, 79°25'’W, 
3050-3150 m: NK _ tape-recordings (12-13 November 1992); 
present study (12-18 February 1996). 


RED-FACED PARROT Hapalopsittaca pyrrhops 


Two pairs with one and two young were observed at Paramos de 


Matanga. There are few known sites for this endangered parrot (Collar 
et al. 1992). 


WHITE-THROATED SCREECH-OWL Otus albogularis 


Recorded at all sites except Mazan and Lagunillas. 


RUFOUS-BANDED OWL Ciccaba albitarsus 


Recorded at all sites except Toledo. 


RUFOUS-BELLIED NIGHTHAWK Lurocalis rufiventris 


Recorded at all sites except Toledo and Salinas. Only doubtfully 


recorded at Mazan. 


N. Krabbe et al. 254 Bull. B.O.C.1997 117(4) 


NEBLINA METALTAIL Metallura odomae | 
Netted at Toledo (3080-3350 m) and Lagunillas (3050-3350 m). 
Previous Ecuadorian records are summarized by Collar et al. (1992). 


There are as yet no specimens to document the reported sympatry of 
M. william with M. baron and M. odomae (Collar et al. 1992). 


MOUNTAIN AVOCETBILL Opisthoprora euryptera 


Found to be fairly common at Oyacachi and Anatenorio, uncommon 
at ‘Toledo. Previously considered rare to uncommon throughout eastern 
Ecuador (Ridgely 1980, own obs., R. S. Ridgely pers. comm.). Rather 
inconspicuous and difficult to record except by its distinctive call or by 
mist-netting or observing near flowers of the genus Centropogon 
(Campanulaceae). Contra Fyjeldsa & Krabbe (1990), the species 
appeared to exploit no other source of nectar. On several occasions it 
was observed to feed on nectar from these flowers by piercing them 
near the. base of the corolla. Its pointed, upturned bill shape may be an 
adaptation to this previously undescribed behaviour. 

Ornelas (1994) has suggested that bill serrations are an adaptation for 
nectar robbing (facilitating ‘illegal’ access through corollas to protected 
nectaries). However, the ZMUC specimen of Opisthoprora euryptera 
does not have serrate tomia despite being a specialised nectar robber 
perforating the base of long-tubed corollas, and the species is not listed 
in Ornelas (1994) as a hummingbird with serrations on the tomia. 
Thus, serrations and pointed/upturned bills are distinct tools which 
may have either different functions (manipulation of insects and 
penetration of corollas, respectively) or a common function (corolla 
piercing). As stated by Ornelas (1994) more evidence is needed from 
ontogenetic, phylogenetic and behavioural studies to support one or the 
other of these views. 


CRESCENT-FACED ANTPITTA Grallaricula lineifrons 


A single bird was called in with tape-recordings of song on three 
occasions at 3320 m at Oyacachi, the type locality. It did not vocalise. 
At Matanga, where it had not been recorded before, we tape-recorded 
songs of it and of the generally lower-elevational Grallaricula nana in 
two adjacent valleys, both at 3085-3150 m, only one species in each 
valley. G. linetfrons was also tape-recorded higher, at .3250m at 
Matanga. No altitude was given for the type specimen (Chapman 1926; 
contra Fjeldsa & Krabbe 1990). All known records of G. linetfrons lie 
between 3000 and 3400 m altitude, in the southern Central Andes of 
Colombia and in the eastern Andes of Ecuador south to Rio Zamora 
(Robbins et al. 1994). 


PALE-FOOTED SWALLOW WNottochelidon flavipes 


Found at Anatenorio, Toledo, and Lagunillas. These records fill 
out important distributional gaps in Ecuador of this overlooked 
cloud-forest swallow, which is probably more or less continuously 
distributed from Santa Cruz Department in Bolivia, through Peru, 
Ecuador and Colombia to Trujillo Province in Venezuela (Parker e¢ al. 
1985, Davis et al. 1994, Ryan & Lentino 1995). 


N. Krabbe et al. 255 Bull. B.O.C. 1997 117(4) 


GREATER SCYTHEBILL Campylorhamphus pucheram 


We observed one of this rare and local species once, at Toledo in 
heavily moss-covered Weinmannia (Cunoniaceae) forest at 3050 m. 


MASKED MOUNTAIN-TANAGER Buthraupis wetmoret 


Found in treeline scrub at Matanga (3250 m), Toledo (3150-3350 m), 
and Lagunillas (3300m). Probably also occurs at Oyacachi and 
Anatenorio, above 3350 m. In 1985, the species was only known in 
Ecuador from one locality (Parker et al. 1985). Recently, it has been 
found in Ecuador at Cerro Mongus, Carchi Province (Robbins ez al. 
1994), Los Llanganates, Napo Province (by NK) and at Cajanuma, 
Loja Province (Bloch et al. 1991). It may thus be distributed 
throughout in Ecuador on the eastern slope in suitable treeline habitat. 


WHITE-CAPPED TANAGER Sericossypha albocristata 


Found at Oyacachi. There are few records from northern Ecuador. 
Also recorded at Matanga and Toledo. 


Acknowledgements 


Our investigation was funded by the Center for Research on Cultural and Biological 
Diversity of Andean Rainforests (DIVA), an interdisciplinary project of collaboration 
between University institutions of Ecuador, Peru, Bolivia and Denmark. We thank MAG 
(Ministerio de Agricultura y Ganaderia, Quito) for permission to undertake research in 
Ecuador and Pedro of MAG in Loja for important advice on Cerro Toledo. 


References: 

Berlepsch, H. & Taczanowski, L. 1884. Deuxiéme liste des oiseaux recueillis dans 
lEquateur donnés au Muséum par M. Clavery. Bull. Mus. natn. Hist. nat. Paris 34: 
71-78. 

Bloch, H., Poulsen, M. K., Rahbek, C. & Rasmussen, J. F. 1991. A survey of the 
montane forest avifauna of the Loja Province, southern Ecuador. ICBP Study 
Report No. 49, Cambridge. 

Chapman, F. 1926. The distribution of bird-life in Ecuador. Bull. Am. Mus. Nat. Hist. 
= i 


Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Parker, T. A. & Wege, 
D. C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data Book. 
International Council. for Bird Preservation, Cambridge, U.K. 

Davis, S. E., Rocha O., O., Sarmiento, J. & Hanagarth, W. 1994. New departmental 
records and notes for some Bolivian birds. Bull. Brit. Orn. Cl. 114: 73-85. 

Fjeldsa, J. & Krabbe, N. 1986. Some range extensions and other unusual records of 
Andean birds. Bull. Brit. Orn. Cl. 106: 115-124. 

Fjeldsa, J. & Krabbe, N. 1990. Birds of the High Andes. Zoological Museum, University 
of Copenhagen and Apollo Books, Svendborg, Denmark. 

Hilty, S. L. & Brown, W. L. 1986. A Guide to the Birds of Colombia. Princeton Univ. 
Press. 

Krabbe, N. 1991. Avifauna of the temperate zone of the Ecuadorian Andes. Zoological 
Museum, University of Copenhagen. Technical report. 

Krabbe, N. 1992. Notes on distribution and natural history of some poorly known 
Ecuadorian birds. Bull. Brit. Orn. Cl. 112: 169-174. 

Meyer de Schauensee, R. & Phelps, W. H. 1978. A Guide to the Birds of Venezuela. 
Princeton Univ. Press. 


Ornelas, J. F. 1994. Serrate tomia: an adaptation for nectar robbing in hummingbirds? 
Auk 111: 703-710. 


N. Krabbe et al. 256 Bull. B.O.C. 1997 117(4) 


Parker, T. A., Schulenberg, T.S., Graves, G. R. & Braun, M. J. 1985. The avifauna of 
the Huancabamba region, northern Peru. Pp. 169-197 im P. A. Buckley et al. (eds). 
Neotropical Ornithology. Orn. Monogr. No. 36, American Ornithologists’ Union. 

Pearman, M. 1994. Neotropical Notebook, Ecuador (Imperial Snipe). Cotinga 1: 26-29. 

Petersson, L. 1996. Neotropical Notebook, Ecuador (Imperial Snipe). Cotinga 5: 78. 

Poulsen, B. O. 1993. Change in mobility among crepuscular ground-living birds in an 
Ecuadorian cloud forest during overcast and rainy weather. Ornitologia Neotropical 
4: 103-105. 

Remsen, J. V. & Graves, W. 5S. 1995. Distribution patterns of Buarremon brush-finches 
(Emberizinae) and interspecific competition in Andean birds. Auk 112: 225-236. 

Ridgely, R. S. 1980. Notes on some rare or previously unrecorded birds in Ecuador. 
American Birds 34: 242-248. 

Robbins, M. B., Ridgely, R. S., Schulenberg, T. S. & Gill, F. B. 1987. The avifauna of 
the Cordillera de Cutuctu, Ecuador, with comparisons to other Andean localities. 
Proc. Acad. Nat. Sci. Philadelphia 139: 243-259. 

Robbins, M. B., Krabbe, N., Rosenberg, G. H., Ridgely, R. S. & Sornoza Molina, F. 
1994. Notes on the natural history of the Crescent-faced Antpitta. Wilson Bull. 106: 
169-173. 

Robbins, M. B., Krabbe, N., Rosenberg, G. H. & Sornoza Molina, F. 1994. The tree line 
avifauna at Cerro Mongus, Prov. Carchi, northeastern Ecuador. Proc. Acad. Nat. 
Sci. Philadelphia 145: 209-216. 

Robinson, F. (ed.) 1987. Rio Mazan Project. 1986 report. Department of Plant Sciences, 
University of Oxford. Unpublished. 

Ryan, T. P. & Lentino R., M. 1995. An additional record of the Pale-footed Swallow 
Notiochelidon flavipes from Venezuela. Bull. Brit. Orn. Cl. 115: 28-29. 

Salvadori, T. & Festa, E. 1899. Viaggio del Dr. Enrico Festa nell’ Ecuador. Uccelli. 
Part II. Passeres Clamatores. Boll. Muset Zool. Anat. comp. R. Umv. Torino 362 
(XV): 1-34. 

Terborgh, J. & Weske, J. S. 1972. Rediscovery of the Imperial Snipe in Peru. Auk 89: 
497-505. 


Addresses: Niels Krabbe, Bent Otto Poulsen and Amy Frelander, Zoological Museum, 
University of Copenhagen, Universitetsparken 15, DK-2100, Copenhagen, 
Denmark. Present address (NK): Casilla 17-21-791, Quito, Ecuador. Present address 
(BOP): The Teachers’ University (DLH), Institute of Biology, Eindropvej 101, 
2400 Copenhagen NV, Denmark. Orfa Rodriguez Barahona, CECIA (Cooperacion 
Ornitologica del Ecuador), Casilla 17-17-906, Quito, Ecuador. 


© British Ornithologists’ Club 1997 


E. P. Toyne & F. N. M. Flanagan 257 Bull. B.O.C. 1997 117(4) 


Observations on the breeding, diet and 
behaviour of the Red-faced Parrot 
Hapalopsittaca pyrrhops in southern Ecuador 


by E. P. Toyne & F. N. M. Flanagan 
Received 16 August 1996 


The Red-faced Parrot Hapalopsittaca pyrrhops is endemic to southern 
Ecuador and immediately adjacent Peru where it is confined to the 
temperate Andean forests between 2300 and 3500m (Collar et al. 
1992). It is regarded as “‘critically endangered’’ (Lambert et al. 1993) 
and listed with other birds “for which their situation was serious and 
action is urgent” (Collar et al. 1992). Red-faced Parrot was previously 
dealt with as a subspecies of the Rusty-faced Parrot Hapalopsittaca 
amazonina but strong evidence to treat it as a full species has now been 
provided (Graves & Uribe Restrepo 1989). Prior to this study virtually 
nothing was known about the biology of this parrot or other members 
of the genus (Forshaw 1989, King 1989, Fjyeldsa & Krabbe 1990, 
Rasmussen et al. 1996). Here we present data on nesting chronology, 
diet and behaviour; information on nesting and vocalizations is 
presented elsewhere (Toyne et al. 1995, Toyne & Flanagan 1996). 


Study area and methods 


The forests of Loja Province were surveyed as part of the series of 
Imperial College “Parrots in Peril’ expeditions which documented the 
status, distribution and biology of parrots in southern Ecuador (Toyne 
1996). Suitable habitat for Red-faced Parrots was surveyed in Loja 
Province in two areas: around Saraguro (3°37'S, 79°14’W) and in 
Podocarpus National Park (PNP) (4°06’S, 79°09'W). Fieldwork was 
conducted in March—May 1992 and October 1994—February 1995. In 
addition, J.N.M.F. visited the Saraguro area intermittently between 
March 1995 and December 1996. For details of localities and habitat 
descriptions see Toyne et al. (1995) and Rasmussen et al. (1996). 
Data on breeding biology were collected at a nest-site near Saraguro 
(Toyne & Flanagan 1996), which was visited between 10 November 
1994 and 8 February 1995, and from general observations through- 
out the study. Behavioural data were collected at the nest-site 
during different stages of breeding. ‘The nest was monitored during the 
incubation period over seven days between 10 November and 
1 December 1994 for approximately 49 hours. On three days (11-12 
and 30 November) the nest was watched from dawn to dusk. During 
the nestling period the nest-site was visited on six days and 
observational data were collected on four days (30 December 1994, 3-4 
and 14 January 1995). On these days a total of approximately 41 hours 
was spent watching the nest. Nestlings were measured using standard 
techniques (Spencer 1984). Elsewhere, when parrots were located, 
basic ecological data such as flock size and food sources were collected 


E. P. Toyne & F. N. M. Flanagan 258 Bull. B.O.C. 1997 117(4) 


(see Toyne et al. 1992 for methods). Specimens of the food plants were 
collected under licence and deposited at the National Herbarium of the 
Museo Ecuatoriano de Ciencias Naturales, Quito; the herbarium of 
Universidad Nacional de Loja, Loja; the Royal Botanical Gardens, 
Kew; and Missouri Botanical Gardens, U.S.A. 


Results 


Flock size 

Red-faced Parrots were encountered in pairs throughout the study 
period and also in flocks of up to 19 individuals. In April and May 
flocks were large (up to 18 individuals) and it was presumed that they 
contained family groups as they comprised adults and newly fledged 
young. In November and December flocks were mostly between four 
and ten. A flock of 19 individuals in November 1994, at Selva Alegre 
near Saraguro, was the largest recorded, but the age class composition 
of this flock was not known. Flocks were usually observed flying above 
the tree canopy (30 m from the ground), or low (1—2 m) on short flights 
(less than 200m) over open ground between wooded habitats. We 
rarely saw them flying very high (>60m from the ground) as some 
other parrots do. Occasionally they perched on isolated trees in open 
land. 

During the survey their numbers appeared to be stable with similar 
sized flocks encountered in the same areas in 1992 and 1994—96. 


Timing of breeding season 

‘The Red-faced Parrot’s breeding season in Loja Province appeared 
to be between October and January. These dates were based on 
monitoring one successful nest (Toyne & Flanagan 1996), observations 
of nest-prospecting pairs in the same area during November and 
December, and sightings of juveniles (identified by plumage 
differences) during January, April and May in three different forests. 

The October—April period, which includes the dry season, is 
probably the breeding season for many bird species in this area, 
as suggested by the following records: Red-backed Hawk Buteo 
polyosoma—with young; Speckle-faced Parrot Pionus tumultuosus— 
nesting; Streaked Tuftedcheek Pseudocolaptes boissonneautii—nesting; 
Cinnamon Flycatcher Pyrrhomyias cinnamomea—nesting; ‘Turquoise 
Jay Cyanolyca turcosa—nest-building; Great Thrush Turdus fuscater— 
pair copulating; Rufous-collared Sparrow Zonotrichia capensis— 
courtship feeding; Mountain Cacique Cacicus leucorhamphus—nesting. 

Red-faced Parrots nested in habitat they used between October and 
June (no surveys have been conducted between July and September), 
suggesting that they were sedentary. Evidence from successful breeding 
in a cavity of a lauracous tree (Toyne & Flanagan 1996) and from three 
pairs prospecting cavities in four trees c. 500 m from this nest indicated 
that they are cavity nesters like the vast majority of parrot species. 
Three of the four cavities were natural holes in Styrax sp. 
(Styracaceae), Symplocos sp. (Symplocaceae) and an unidentified tree. 
The fourth hole looked like a previously used nest-hole in an 


E. P. Toyne & F. N. M. Flanagan 259 Bull. B.O.C. 1997 117(4) 


unidentified tree. On two occasions flocks of six parrots were observed 
visiting these holes in November and December. These flocks split into 
pairs, each of which visited different cavities. On one occasion two 
individuals from different pairs fought to enter a hole. Despite several 
checks none of the four cavities was used between November and 
January. One bird, which visited a Streaked ‘Tuftedcheek’s cavity nest 
in a tree, was presumed to be nest-prospecting. 

The approximate nesting chronology of Red-faced Parrots could 
be estimated from observations at the successful nest. On 10 
November, when the nest was discovered, the hen spent several hours 
in the nest but also left the nest for longer periods than we observed 
later, when she certainly was incubating. We suspect the hen was either 
laying or had just started incubation. On 21 November 1994 the nest 
tree was climbed and the nest (17.5 m above the ground) was inspected, 
and contained two creamy matt white, ovoid eggs. When the nest was 
next inspected on 10 December 1994, it contained two featherless 
nestlings, covered in grey down and with closed eyes. They were 
estimated to be 2-4 days old +1 day. On 3 January 1995 the nestlings 
were re-examined; on this date one was noticeably larger than its 
sibling (see Toyne & Flanagan 1996) and both were aged at 27 + 2 days. 
This age was later validated by aviculturists independently from 
photographs (T. Arndt im litt. 1995). On 14 January 1995 both 
nestlings (c. 38 days old) were seen at the nest entrance. They still had 
down on their wings and backs, and their heads (crown, forehead and 
chin/throat) were strongly marked red. In this they differed from 
adults, which have a green throat and crown with red forehead and 
lores. By 25 January 1995 both young had fledged and were seen 
around the nest. Juveniles, some of which were fed by adults, were 
recorded in PNP and at two woods near Saraguro in April 1992. 
Juveniles were also observed in the same wood as the nest between 9 
and 11 May 1989 (Rasmussen et al. 1996) and in January 1994. These 
data give an idea of the nesting chronology: pre-laying period, October; 
egg-laying and incubation period (c. 26-29 days), November-— 
December; nestling period (c. 49-52 days), December—January; 
post- -fledging period, January—May. 

During October—December 1995 and November—December 1996 the 
nest was not re-used, although Red-faced Parrots were seen around 
the nest tree. 


Behaviour of adults during incubation and nestling period 

As both adults were of similar plumage it was not possible to tell for 
certain if the pair shared incubation and care of nestlings. However, the 
behaviour of the non-incubating bird, presumed to be male, was similar 
throughout the incubation and nestling period. So we assumed the 
female incubated and brooded the young, as is usual for most parrots 
(Forshaw 1989). 

‘The average time a bird spent incubating during daylight hours, i.e. 
between returning to the nest and leaving it, was 192 minutes (n=7, 
range 115-290, s.d. 59.2). At feeding time one parrot approached the 
nest-site, calling. The incubating bird would then climb to the entrance 


E. P. Toyne & 7. N. M. Flanagan 260 Bull. B.O.C. 1997 117(4) 


of the nest and after 1 or 2 minutes they would fly off silently together. 
They flew 20 m to a mature lauraceous tree where the non-incubating 
bird would feed its partner with regurgitated food. The parrots would 
leave the nest unattended for an average of 9 minutes (n=11, range 
1-21, s.d. 7.8). The incubating bird was fed three times a day, gener- 
ally between 07.00—08.30, 12.00-13.00 and 15.00-17.00 hours. The 
incubating bird was nearly always called off the nest by its mate and 
flew to a neighbouring tree to be fed by it; only once was it fed at the 
nest entrance. 

The nestlings when c. 23+days old were not brooded but left 
unattended throughout most of the day until one adult returned to the 
nest, presumably to feed them. The average time between feeds was 
113 minutes (n=13, range 14—250, s.d. 90.2). Adults fed the nestlings 
with regurgitated food on average five times a day. The adults spent 
little time at the nest (5.8 minutes, n=12, range 2-11, s.d. 2.9). The 
nestlings were brooded at night by an adult who entered the nest 
around 18.25 h. 


On 14 January 1995 the young were at least 38 days old and, during 
that day, they were fed on four occasions at the nest entrance. During 
feeds each nestling was fed twice with regurgitated food. The nestlings 
were brooded that night. 


‘The parrots did not appear to defend their nest against intruders; at 
any rate they showed no alarm when Mountain Caciques or Turquoise 
Jays landed on the nest tree and perched near or on the nest entrance. 
On 14 January 1995 a Strong-billed Woodcreeper (Xzphocolaptes 
promeropirhynchus) on two separate occasions landed at the nest 
entrance and looked into the nest but did not enter; both adult parrots 
were absent from the nest. Once a pair of Red-faced Parrots visited the 
nest tree and one of them entered the nest whilst the nest was occupied 
by a brooding bird and the two nestlings. The intruder left, presumably 
because it found the nest occupied. 


Roosting habitat and roosting behaviour 

On the evenings of 12 and 14 April 1992 Red-faced Parrots roosted 
in the 1994 nest tree. Although this tree was in fruit, no parrots were 
seen to feed from it. Roosting numbers increased each day; 4 on day 
one, 7 on day two and 18 on day three (observations then had to cease). 
In the morning (06.00-07.00 h) they left the roost and settled in 
adjacent trees before finally dispersing in small groups of two to six in 
all directions. The parrots arrived at the tree in small flocks (3-7) 
between 17.30 and 18.50 h. Before landing in the roost tree they would 
fly around the perimeter of the field, landing in the adjacent trees and 
calling loudly. While doing so, they would be joined by late-comers. 
On 14 April, when 18 birds came to roost, some flew into a nearby tree 
before entering the roost tree, whilst others flew directly into the top 
quarter of the roost tree. On all evenings, when the parrots were 
settling down to roost one parrot would perch on the crown of the tree 
in a look-out position and generally wait for 10 minutes before 
disappearing to roost with the rest of the flock. 


E. P. Toyne & }. N. M. Flanagan 261 Bull. B.O.C. 1997 117(4) 


Feeding ecology 

Red-faced Parrots were recorded feeding on the following plants, all 
except Myrcianthes sp. in the Saraguro area. The only food previously 
recorded is Phytolacca berries (Fjeldsa & Krabbe 1990). 

Vriesea sp. (Bromeliaceae). The white stamens within the terminal 
inflorescence of these epiphytic plants were eaten in February 1995. 
The flowers were not open and the parrots had to remove the exterior 
petals with their bills to reach the stamens. In December one H. 
pyrrhops was observed drinking water that had collected 1n the leaves of 
an epiphytic bromeliad. 

Virburnum leptophyllum (Caprifoliaceae). The berries of this tree 
were eaten in April 1992. 

Weinmannia latifolia and Weinmannia pinnata (Cunoniaceae). The 
shoots, flowers and seeds of these trees were eaten in April 1992. One 
adult fed W. latifolia to a begging juvenile (‘Toyne et al. 1995). 

Weinmannia elliptica (Cunoniaceae). The flowering parts and shoots 
from four trees of this species were eaten in late November 1994 and 
January 1995. In the latter month adults were observed feeding 
juveniles with this food. 

Miconia jahnii (Melastomataceae). The berries of this tall tree were 
eaten in April 1992. 

Miconia sp. (Melastomataceae). The green berries of this tall tree 
were eaten in November 1994 and February 1995. 

Miconia sp., or Calyptrella stellata (Melastomataceae). The yellow 
seed pods were eaten throughout November and January (1994/95). 

Myrcianthes rhopaloides (Myrtaceae). ‘The seed pods of this tree were 
eaten during November and December in 1994 and 1996. 

Myrcianthes sp. (Myrtaceae). The flowers of this tree were eaten at 
Cajanuma (PNP) in January 1995. 

Aegiphila sp. (Verbenaceae). ‘The dark green seed pods of this tree 
were eaten in November 1994 and February 1995. 

Clethra revoluta (Clethraceae). The white flowers and seed pods were 
eaten in April 1992. 

Cavendishia bracteata (Ericaceae). The berries from this 8m tall 
plant growing in secondary forest were eaten in May 1992. 

Disterigma alaternoides (Ericaceae). The berries from this plant, a 
scrambling 5m tall shrub in secondary growth forest, were eaten in 
May 1992. 

On 24 November 1996 J.N.M.F. observed a flock of 13 land on the 
ground next to the base of a tree stump. The ground there consisted of 
grass and bare soil. One parrot perched on the stump whilst the others 
foraged on ground that was selected as it was in the shade and provided 
some cover from predators. The look-out parrot changed several times. 
This observation is interesting as it probably represents the only case of 
this species foraging on soil, presumably for minerals. 


Discussion 


Clearly, one cannot draw many conclusions about the Red-faced 
Parrot’s breeding biology from observations of one nesting attempt. 


E. P. Toyne & F. N. M. Flanagan 262 Bull. B.O.C. 1997 117(4) 


Due to the lack of data on Hapalopsittica species our data can only be 
compared to other similar sized parrots such as Pionopsitta species 
(Forshaw 1989). The Pileated Parrot Pzionopsitta pileata has an 
incubation period of 24 days (cf. Red-faced Parrot: 26-29 days), 
fledging between 52-54 days of age (cf. Red-faced Parrot: 49-52 days) 
and self-feeding when 57-59 days old (Forshaw 1989). These data 
suggest that the Red-faced Parrot’s incubation and fledging periods are 
similar to those of other parrots of similar size. 

The fact that Red-faced Parrots feed on common Andean plant 
genera such as Micoma, Myrcianthes, Weinmannia and Clethra suggests 
that their restricted range and their rarity are not due to a dependence 
on certain foods. However, they may require special plants at certain 
times such as their breeding season. For example in October, the 
pre-laying period of most birds in the yungas of Cochabamba, Bolivia, 
Black-winged Parrots Hapalopsittaca melanotis were found to specialise 
on the fruits of cloud-forest mistletoes of the genus Gatadendron, which 
may represent a high-nutrition food source (J. Fjeldsa zn litt. 1993). 


Acknowledgements 


The following people participated in the fieldwork: Pablo Andrade, Mark Jeficote, 
Adrian Griffiths, Gabi Howard, Sachin Kapila, Orfa Rodriguez, Rodrigo Tapia, and 
Tracy-Ann Hooley. The authors would like to thank INEFAN in Quito, Loja and 
Zamora for permission to work in Ecuador, and the Saraguro Indian communities 
and various landowners for permission to visit their land. The Ecuadorian Embassy in 
London and the British Embassy in Quito provided advice and guidance. Corporacion 
Ornitologica del Ecuador (CECIA) are thanked for logistical support in Ecuador, as are 
Fundacion Ecologica Arcoiris and Promusta for their help whilst the expedition was in 
Loja Province. The following botanists helped identify parrot food sources: David Neill, 
Terry Pennington, John Wurdack, Claes Gustafsson and Bolivar Merino. Thomas 
Arndt, Nigel Collar, Jon Fyjeldsa, Bent Otto Poulsen, Robert Ridgely, Thomas 
Schulenberg and an anonymous referee kindly commented on a previous draft. Lastly, 
we wish to acknowledge our sponsors who made the expeditions possible: Imperial 
College Exploration Board, Imperial College Biology Department, Royal Geographical 
Society, British Ornithologists’ Union, British Ecological Society, BirdLife 
International/Fauna and Flora International BP conservation award, Bird Exploration 
Fund, Frederick Gregory Fund, Gilchrist Educational Trust, Mount Everest 
Foundation, The World Parrot Trust, The Parrot Society of the U.K., Barclays Bank, 
British Petroleum, River Island Clothing Company, Rob Thompson Memorial Fund, 
Journey Latin America, East Productions Ltd. and Wildwings. 


References: 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. J., Parker, 
T. A. & Wege, D. C. 1992. Threatened Birds of the Americas: the ICBP/IUCN Red 
Data Book. ICBP, Cambridge, U.K. 

Fjeldsa, J. & Krabbe, N. 1990. Birds of the High Andes. Zoological Museum, University 
of Copenhagen and Apollo books, Svendborg, Denmark. 

Forshaw, J. M. 1989. Parrots of the World (3rd edn). Blandford Press, U.K. 

Graves, G. R. & Uribe Restrepo, D. 1989. A new allopatric taxon in the Hapalopsittaca 
amazonina (Psittacidae) superspecies from Colombia. Wilson Bull. 101: 369-376. 

King, J. R. 1989. Notes on the birds of the Rio Mazan Valley, Azuay Province, Ecuador; 
with special reference to Leptosittaca branicku, Hapalopsittaca pyrrhops and 
Metallura baron. Bull. Brit. Orn. Cl. 190: 140-147. 

Lambert, F., Wirth, R., Seal, U. S., Thomsen, J. B. & Ellis-Joseph, S. 1993. Parrots: An 
action plan for their conservation 1993-1998. Unpublished second draft. IUCN and 
BirdLife International, Cambridge, U.K. 


E. P. Toyne & 7. N. M. Flanagan 263 Bull. B.O.C. 1997 117(4) 


Rasmussen, J. F., Rahbek, C., Poulsen, B. O., Poulsen, M. K. & Bloch, H. 1996. 
Distributional records and natural history notes on threatened and little known birds 
of southern Ecuador. Bull. Brit. Orn. Cl. 116: 26-46. 

Spencer, R. 1984. The Ringers’ Manual (3rd edn). B.T.O., Norfolk, U.K. 

Toyne, E. P. 1996. The Imperial College Parrots in Peril initiative: a summary of the 
1990, 1992 & 1994 expeditions. Bull. B.E.S. 17: 222-224. 

Toyne, E. P. & Flanagan, J. N. M. 1996. First nest record of Red-faced Parrot 
Hapalopsittaca pyrrhops in Ecuador. Cotinga 5: 43-45. 

Toyne, E. P., Flanagan, J. N. M. & Jeffcote, M. T. 1995. Vocalizations of the endangered 
Red-faced Parrot Hapalopsittaca pyrrhops in southern Ecuador. Ornitologia 
Neotropical 6: 125-128. 

Toyne, E. P., Jeffcote, M. T. & Flanagan, J. N. M. 1992. Status, distribution and ecology 
of the White-breasted Parakeet Pyrrhura albipectus in Podocarpus National Park, 
southern Ecuador. Bird Conserv. Int. 2: 327-339. 


Addresses: E. P. Toyne, Department of Biology, Imperial College of Science, Technology 
and Medicine, London SW7 2BB, U.K. Current address: WWF-UK, Panda House, 
Weyside Park, Cattershill Lane, Godalming, Surrey GU7 1XR. J. N. M. Flanagan, 
Fundacion Ecologica Arcoiris, Casilla 11-01-860, Loja, Ecuador. 


© British Ornithologists’ Club 1997 


P.W. Smith 264 Bull. B.O.C. 1997 117(4) 


The history and taxonomic status of the 
Hispaniolan Crossbill Loxia megaplaga 


by P. William Smith 


Received 5 October 1996 


In October 1916, W. L. Abbott collected a pair of hitherto unknown 
and unexpected crossbills near El Rio, in the Cordillera Central of the 
Dominican Republic, on the West Indian island of Hispaniola. These 
were described by Riley (1916) as Loxia megaplaga (holotype USNM 
249515), and were compared in plumage and morphology primarily 
with the palearctic L. bifasciata, then generally considered a species 
distinct from the nearctic L. leucoptera, the White-winged Crossbill 
(but see Hartert 1910). In his comments, Riley asserted that megaplaga 
‘‘can be told at a glance’’ from leucoptera by its heavier, less attenuated 
bill, a difference further discussed and illustrated by Richmond (1916). 
As he was leaving Hispaniola, Abbott encountered Rollo Beck, then 
collecting for the Brewster-Sanford Collection. Told by Abbott of this 
and other interesting discoveries, Beck spent much of late February and 
March 1917 in the Cordillera Central (Beck 1921), eventually securing 
a series of thirty-one specimens of megaplaga, including several recent 
fledglings. ‘Twenty-three of these are now at the American Museum of 
Natural History and four are at the British Museum (Natural History). 
In a search of likely museums I was able to learn of less than ten 
specimens taken subsequently, suggesting that fewer than fifty exist. 
When Beck’s series came to the attention of Frank Chapman at the 
American Museum of Natural History, Chapman described the 
discovery as ‘‘one of the ornithological sensations of recent years”’ 
(Chapman 1917). After saying that he had no specimens of bifasciata 
for comparison, he speculated how a “‘race’? (Chapman’s word, 
although he maintained binomial nomenclature and referred to it as a 
““species’” elsewhere in his note) of the North American White-winged 
Crossbill leucoptera (s.s.) could occur in such a remote, tropical 
environment. He suggested that White-winged Crossbills, as well as 
the pines to which they were adapted (sic, infra), may have been more 
widespread during the previous ice age. Neither Bond (1928), Wetmore 
& Swales (1931), nor the A.O.U. (1931) considered megaplaga (or 
bifasciata) to be a subspecies of leucoptera, although Wetmore & Swales 
(1931) stated that ‘““Obviously the three are from common stock’’. 
The first explicit treatment of megaplaga as a subspecies of leucoptera 
(s.l.) seems to have been by Hartert & Steinbacher (1932). Bond (1936) 
also used trinomial treatment, as did Hellmayr (1938), although neither 
cited Hartert & Steinbacher (1932); Griscom (1937), on the other hand, 
did so. The A.O.U. formally adopted subspecific treatment for 
megaplaga several years later (A.O.U. 1944), citing Hellmayr (1938). It 
is not evident that any of those authors were expressing other than the 
emergent taxonomic philosophy of the era, which focused more on 
apparent similarities between taxa than differences. Bond (1945) wrote, 


P. W. Smith 265 Bull. B.O.C. 1997 117(4) 


‘The presence of a White-winged Crossbill in Hispaniola is probably 
the result of an invasion of the species during the Pleistocene’, a notion 
for which Chapman (1917) apparently laid ground and which Wetmore 
& Swales (1931) had developed further. 

Griscom (1937) is the principal author to have analysed the 
alignment of megaplaga with leucoptera, his discussion also including 
bifasciata. He rejected six of nine possible criteria proposed by other 
authors to diagnose L. leocoptera (s.l.) from L. curvirostra (s.l.) as 
overlapping: smaller size and bill; proportionately more slender bill; 
pinker or paler shades of red in adult males; yellower olive tones in 
adult females; two well-developed white wing-bands (!); and different 
breeding and winter plumage of adult males. He accepted only broad 
white tips to the tertials, blackish remiges and rectrices, and black 
upper tail coverts as characters shared by all forms of leucoptera and no 
forms of curvirostra. 

Meanwhile, megaplaga largely slipped from ornithological conscious- 
ness. It apparently went unreported in field literature for the forty years 
following Bond’s collecting five in Haiti in 1930 (unpublished, M. 
Robbins 77 /itt.). Annabelle Stockton Dod then claimed to ‘‘rediscover’’ 
crossbills on Hispaniola in 1971 in the Sierra de Baoruco, an extension 
of the Haitian Massif de la Selle into the southwestern Dominican 
Republic (Dod 1978). In fact, J. W. Terborgh had reported them both 
there and in the Cordillera Central a year earlier (Bond 1971). In April 
1971, Dod and the Keplers found the first nest of megaplaga recorded, 
near the Dominican-Haitian border in those same mountains (Kepler 
et al. 1975). At least three crossbills identified carefully and credibly as 
megaplaga were observed in introduced pines in the Blue Mountains of 
Jamaica for several months beginning in December 1970 (Lack et al. 
1972, notwithstanding Bond’s 1972 disbelief), where no species of pine 
is native. There apparently are no other known reports of crossbills in 
Jamaica (R. Sutton) or elsewhere in the West Indies. 

Since its “‘rediscovery’’, megaplaga has been reported occasionally in 
the Dominican Republic by resident and visiting birdwatchers and 
scientists, both in the Cordillera Central and the Sierra de Baoruco. 
Because of near total deforestation, its current status in Haiti is 
uncertain, although apparently it is not completely extirpated there 
(Benkman 1994). Benkman (1994) observed and studied its morphology 
and ecology in the Dominican Republic and concluded that it was a 
specialist adapted to forage solely on Pinus occidentalis, Hispaniola’s 
sole native conifer, found only there and in the Sierra Maestra of 
Cuba (Sauget & Liogier 1946, Mirov 1967). He suggested that its 
bill structure was so different from Jleucoptera (s.s.), primarily a 
spruce-feeding specialist (Benkman 1992), that hybrids would be 
selectively disadvantaged (Benkman 1994). Benkman (1994) computed 
a likely population size for megaplaga of less than 1000 remaining 
individuals. 

In March, 1996, assisted by tape playback of vocalizations which 
Reynard (1981) secured both in Haiti and near El] Aguacate, Dominican 
Republic (G. Reynard im IUitt.), L. Manfredi and I searched 
unsuccessfully for crossbills for five days between c. 700 and 1200 m in 


P. W. Smith 266 Bull. B.O.C. 1997 117(4) 


the Cordillera Central within c. 50 km of Jarabacoa, including the area 
of El Rio where most of Beck’s specimens had been taken. Although 
there still are many patches of pines in that region, few remaining trees 
appear to consist of old growth. An intensive two-year study in a 
portion of the area we surveyed failed to encounter the species (Latta & 
Wunderle 1996), although ornithologists working in the area’s pines 
occasionally do encounter crossbills (Benkman 1994, J. Wunderle). 
Then moving to the Sierra de Baoruco on 26 March to the vicinity of 
the abandoned Zapotén logging camp, 5 km south of El Aguacate on 
the Haitian border at c. 1500m, we attracted a family group of 
crossbills (male, female, and three juveniles) to our location within a 
few minutes of our arrival in an area of old growth by playing the 
Reynard (1981) recording. 

We made leisurely telescopic studies for about an hour between 
8—9 a.m. Our most unexpected observation in light of Griscom’s (1937) 
diagnosis of leucoptera (s.l.) including megaplaga, was that the remiges, 
rectrices, and upper tail coverts did not appear “‘blackish’’ but instead 
appeared brown, and the tertials of the adults did not seem at all 
white-tipped (those on the juveniles were narrowly edged with white). 
Other characters noted included the massive beaks and the fact that the 
greater covert tips seemed only narrowly white, narrower than the 
median covert tips. The birds seemed remarkably silent for crossbills, 
although the male did warble faintly for lengthy periods as we watched. 
Their principal vocalizations, made occasionally during their slow 
approach to our vicinity as we played the recording, were aurally 
indistinguishable to us from those published by Reyard (1981). 

I examined the series of twenty-three specimens of megaplaga, all 
secured by Beck, at the American Museum of Natural History. Susan 
A. Smith and I took measurements of most adults (Table 1), and to the 
extent specimens were available, we also took equivalent measurements 
of a comparable number of adult leucoptera (s.s.), bifasciata, and Loxia 
curvirostra mesamericana, the geographically nearest member of Loxza, 
which is found primarily in pine forest from Guatemala to Nicaragua 
(Howell in Paynter 1968). Additionally, through the courtesy of Brian 
Nelson, I obtained sonagrams of megaplaga’s basic vocalizations from 
Reynard (1981) (Fig. 1). 

In comparing the series of megaplaga with those of other crossbill 
taxa, I failed to discern that megaplaga (or bifasciata) had blackish 
remiges, rectrices, or upper tail coverts, contra Griscom (1937). My 
judgement was that all Loxia have those feathers concolorous. 
Leucoptera’s (s.s.) indeed are black, but all other taxa are shades of 
brown, those of bifasciata appearing hardly darker than most 
curvirostra’s and lighter than some. Megaplaga’s feathers are a darker 
shade of brown than bifasciata’s. Adult megaplaga in series also lack 
prominent white tertial tips as shown by many leucoptera (s.s.) and 
bifasciata. | thus concluded that the specimens of megaplaga generally 
agreed with my field observations, but not with Griscom’s ae 
museum basis for including megaplaga within leucoptera (s.1/.), 
discrepancy I cannot explain inasmuch as the same material was largely 
used. 


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Figure 1. Hispaniolan Crossbill vocalizations (from Reynard 1981). 


Unlike either leucoptera (s.s.) or bifasciata, megaplaga has relatively 
narrow white tips to the greater coverts (Table 2), with most such 
feathers being medially dark almost to the edge. This character agrees 
more with the rare, so-called ‘rubrifasciata’ morph of curvirostra (Eck 
1981, Cramp & Perrins 1994). But for the presence of wingbars, the 
only morphological character of megaplaga that suggests commonality 
with leucoptera (s.l.) is the wing:tail ratio (Table 2, Eck 1981). Even that 
feature is Somewhat ambiguous, however, for megaplaga’s ratio is larger 
than either leucoptera’s (s.s.) or bifasciata’s (nearer the latter than the 
former) and begins to approach the various forms of curvirostra (s.l.) 
(Table 2, Eck 1981, Cramp & Perrins 1994). 

The vocalizations of megaplaga (Fig. 1) consist mainly of a series of 
somewhat harmonic 100 ms notes c. 1.2 kHz in range, centred at c. 
3.1 kHz and uttered at a rate of c. 5.5/sec. Among the several calls of 
leucoptera (s.s.) documented by Mundinger (1979), the ‘‘chutter”’ 
consists of notes structurally similar to those of megaplaga but of about 
one-fourth the duration delivered about four times as fast. All other 
calls recorded by Mundinger (1979) are more different structurally. 
The song of leucoptera (s.s.) shown in Cramp & Perrins (1994) consists 
mainly of similar-structured notes of slightly higher frequency than 
those of megaplaga, delivered about twice as fast. I am uncertain 
whether the vocalizations of megaplaga on Reynard (1981) or heard by 
us should be characterized as “‘song’’ or “‘calls’’. The calls of bzfasciata 
documented in Cramp & Perrins (1994) have more harmonic qualities, 
are longer and are given more slowly than vocalizations of megaplaga, 
but span roughly the same frequency. Mundinger (1979) argues that 
vocalizations of carduelines are learned in a social context, thus one 
would expect those of modern isolated megaplaga to have drifted from 
their progenitor. Although Groth (1993) did not study vocalizations of 
mesamericana, none of the calls of North American curvirostra (s.l.) that 
he did document seem as similar structurally to those of megaplaga as 
were those of either leucoptera (s.s.) or bifasciata. 

Bond (1945) and others supposed that megaplaga was a relict of a 
Pleistocene irruption of leucoptera (s.s.). While one cannot dismiss such 


P. W. Smith 269 Bull. B.O.C. 1997 117(4) 


a theory, no particular evidence supports it. Bond's (1940) suppo- 
sition that megaplaga’s presence on Hispaniola “*... is probably due to 
some accidental invasion in the past, the birds becoming established 
on finding conditions suitable’ failed to recognize leucoptera’s (s.s.) 
adaptation to Picea (Benkman 1992). Megaplaga is the only member 
of leucoptera (s.l.) adapted to feed on large cones of Pinus (Benkman 
1994). Mirov (1967) found chemical similarities between Hispaniolan 
P. occidentalis and various species of pine found in central and western 
North America which are lacking from all pine species found in east- 
ern North America. Thus, P. occidentalis probably evolved from 
mesamerican pine stock. It seems plausible, therefore, that megaplaga 
either evolved in situ or colonized from a Central American ancestor. It 
is striking that megaplaga’s bill is rather similar in size to mesamericana 
(Tables 1, 2), which suggests a common ancestor or convergence due to 
similarity in cone structure of the resident pines. Its ancestor may have 
been unlike modern leucoptera (s.s.), which may have evolved its 
spruce-adapted bill structure after that supposed invasion, given that 
there is no evidence that small-coned conifers ever were present in the 
West Indies. 

In plumage and morphology, megaplaga reflects a mosaic of 
characters found in Loxia. As recognized by its describer (Riley 1916), 
it shows more morphological similarity to the palearctic bifasciata than 
the nearctic leucoptera (s.s.). The colour of its flight feathers and the 
width of the white on its greater covert tips are intermediate between 
leucoptera (s.s.) and curvirostra (s.l.). Its wing:tail ratio is closer to 
leucoptera (s.l.) than to curvirostra (s.l.) but is intermediate and more 
like bifasciata than leucoptera (s.s.). It shows morphological similarity to 
the Central American Red Crossbill L. curvirostra mesamericana but 
with ‘rubrifasciata’-like wingbars and a smaller wing:tail ratio. Its 
vocalizations are more like leucoptera (s.l.) than any population of 
curvirostra (s.l.) for which recordings are available (Mundinger 1979, 
Groth 1993, Cramp & Perrins 1994). 

The evolutionary history of megaplaga can be unravelled with 
certainty only by use of genetic techniques. Nevertheless, it now is so 
isolated spatially, morphologically, and ecologically from Jleucoptera 
(s.s.) (Benkman 1994, this study) that nothing seems served by 
combining it with bifasciata and leucoptera into a composite species 
which is questionably diagnosable (Griscom 1937, this study). I 
therefore agree with Benkman (1994) that Loxia megaplaga should be 
recognized as a full species, the Hispaniolan Crossbill, as it was 
described originally. 

Such recognition might also help bring more attention to this 
evidently endangered taxon. Nothing in my travels in the Dominican 
Republic since 1987 has led me to believe that Benkman’s (1994) 
population estimate of under 1000 individuals was unduly pessimistic. 
Most reports of megaplaga since 1970 have been along the Haitian 
border in the Sierra de Baoruco. Although this region is gazetted as 
National Park, there seems to be no active protection. I observed 
widespread poaching of trees by Haitians, who widely lack their own 
trees, and several active charcoal-making sites along the border in the 


P. W. Smith 270 Bull. B.O.C. 1997 117(4) 


Zapotén area. Without more active protection, the trees to which 
megaplaga is specifically adapted seem likely to disappear, as they 
largely have across the border. So probably would vanish a unique 
avian relict. 


Acknowledgements 


Allison Andors, Mary LeCroy, and the staff of the American Museum of Natural 
History, New York, facilitated my study of crossbill specimens, assisted by Susan A. 
Smith, while Mr & Mrs Kenneth Prytherch kindly hosted a visit there. M. P. Adams 
(British Museum (Natural History), Tring), Ralph Browning (United States National 
Museum, Washington), Karen Cebra (California Academy of Sciences, San Francisco), 
Per Ericson (Swedish Museum of Natural History, Stockholm), Simén Guerrero 
(Museo Nacional de Historia Natural, Santo Domingo), Janet Hinshaw (University of 
Michigan Museum of Zoology, Ann Arbor), James Laughlin (Carnegie Museum 
of Natural History, Pittsburgh), Van Remsen (Louisiana State University Museum of 
Natural Science, Baton Rouge), Mark Robbins (Academy of Natural Sciences, 
Philadelphia), and David Willard (Field Museum of Natural History, Chicago) all 
furnished information about their collections, while Janet Hinshaw and others 
associated with the Josselyn Van Tyne Memorial Library of the Wilson 
Ornithological Society at the University of Michigan secured much helpful literature. 
The Natural Resources Center in Everglades National Park, Florida, provided library 
and other support services. Craig Benkman, John Elmberg, and Jeffrey Groth freely 
discussed their crossbill work with me and also provided useful literature. George 
Reynard kindly allowed me to incorporate sonagrams from his recordings into my 
study, and Brian Nelson of the Archbold Biological Station, Lake Placid, made those 
sonagrams on his personal computer using software from the Cornell Laboratory of 
Ornithology. David Lee, Mark van Biers, Joseph Wunderle, and others provided 
suggestions for finding crossbills on Hispaniola. William Beaty greatly facilitated my 
travels in the Dominican Republic, and he, along with Simon Guerrero, Larry 
Manfredi, Susan A. Smith, Mary Catherine Wheeler, and Robert Whitcomb, all 
provided field assistance and companionship. Craig Benkman, Jeffrey Groth, and 
William Robertson all offered constructive suggestions on earlier drafts of this 
manuscript. Without their combined assistance, I could not have completed this 
study, and I sincerely thank them all. 


References: 

A.O.U. 1931. Check-List of North American Birds. 4th edn. American Ornithologists’ 
Union, Lancaster, Pennsylvania. 

A.O.U. 1944. Nineteenth supplement to the American Ornithologists’ Union check-list 
of North American Birds. Auk 61: 441-464. 

Beck, R. 1921. Bird collecting in the highlands of Santo Domingo. Nat. Hist. 21: 37-49. 

Benkman, C. W. 1992. White-winged Crossbill. No. 27 in The Birds of North America 

. (A. Poole, P. Stettenheim and F. Gill, eds). Academy of Natural Sciences, 
Philadelphia, Pennsylvania. 

Benkman, C. W. 1994. Comments on the ecology and status of the Hispaniolan Crossbill 
(Loxia leucoptera megaplaga), with recommendations for its conservation. Caribbean 
F., Sct. 30: 250-254. 

Bond, J. 1928. The distribution and habitats of the birds of the Republic of Haiti. Proc. 
Acad. Nat. Sci. Philadelphia 80: 483-521. 

Bond, J. 1936. Birds of the West Indies. Academy of Natural Sciences, Philadelphia, 
Pennsylvania. 

Bond, J. 1940. Check-list of birds of the West Indies. Jbid. 

Bond, J. 1945. Check-list of birds of the West Indies. 2nd edn. Ibid. 

Bond, J. 1961. Birds of the West Indies. First American edition. Houghton Mifflin, 
Boston, Massachusetts. 

Bond, J. 1971. Sixteenth supplement to the check-list of birds of the West Indies (1956). 
Academy of Natural Sciences, Philadelphia, Pennsylvania. 

Bond, J. 1972. Seventeenth supplement to the check-list of birds of the West Indies 
(1956). Ibid. 


P.W. Smith 271 Bull. B.O.C. 1997 117(4) 


Chapman, F. M. 1917. Descriptions of new birds from Santo Domingo and remarks 
on others in the Brewster-Sanford Collection. Bull. Am. Mus. Nat. Hist. 37: 
327-334. 

Cramp, S. & Perrins, C. M. (eds). 1994. The Birds of the Western Palearctic. Vol. 8. 
Oxford Univ. Press. 

Dod, A. S. 1978. Aves de la Republica Dominicana. Museo National de Historia Natural, 
Santo Domingo. 

Eck, S. 1980 (1981). Reflexionen tiber die Taxonomie westpalaearktischer Loxia-Arten 
(Aves, Passeriformes, Fringillidae). Zool. Ahb. Mus. Tierk. Dresden 37: 183-207. 
Griscom, L. 1937. A monographic study of the Red Crossbill. Proc. Boston Soc. Nat. 

Hist. 41: 77-210. 

Groth, J. G. 1993. Evolutionary differentiation in morphology, vocalizations, and 
allozymes among nomadic sibling species in the North American: Red Crossbill 
(Loxia curvirostra) complex. Univ. Cal. Pub. Zool. 127. 

Hartert, E. 1910. Die Végel der paldarktischen Fauna. Vol. 1. R. Friedlander und Sohn, 
Berlin. 

Hartert, E. & Steinbacher, F. 1932. Die Végel der paldarktischen Fauna. Erganzungs- 
band, Heft 1. Berlin. 

Hellmayr, C. E. 1938. Catalog of birds of the Americas and the adjacent islands in Field 
Museum of Natural History. Field Mus. Nat. Hist. Zool. Series 13, Part 11. 

Kepler, A. K., Kepler, C. B. & Dod, A. 1975. First nest record of the White-winged 
Crossbill in Hispaniola. Condor 77: 220-221. 

Lack, D., Lack, E., Lack, P. & Lack, A. 1972. Transients in Jamaica 1970-71. Gosse Bird 
Club Broadsheet 18: 1-5. 

Latta, S. C. & Wunderle, J. M., Jr. 1996. The composition and foraging ecology of 
mixed-species flocks in pine forests of Hispaniola. Condor 98: 595-607. 

Muirov, N. T. 1967. The genus Pinus. Ronald Press Co., New York. 

Mundinger, P. C. 1979. Call learning in the Carduelinae: ethological and systematic 
considerations. Syst. Zool. 28: 270-283. 

Paynter, R. A., Jr. (ed.) 1968. Peters’ Check-List of Birds of the World. Vol. 14. Museum 
of Comparative Zoology, Harvard. 

Reynard, G. B. 1981. Bird songs in the Dominican Republic [cassette]. Cornell 
Laboratory of Ornithology, Ithaca, New York. 

Richmond, C. W. 1916. Explorations in Santo Domingo. Smithsonian Misc. Coll. 66(17): 
36-39. : 

Riley, J. H. 1916. Three remarkable new species of birds from Santo Domingo. 
Smithsonian Misc. Coll. 66(15): 1-2. 

Sauget, J. S. & Liogier, E. E. (“Hermano Leon F.S.C.”’ and ‘“‘Hermano Alain F.S.C.’’) 
1946. Flora de Cuba. Parte I. Contrib. Occ. Mus. Hist. Nat. Colegio de la Salle 
(Habana), Num. 8. Reprinted 1974 by Otto Koeltz Science Publishers, Koenigstein, 
Germany. 

Wetmore, A. & Swales, B. H. 1931. The birds of Haiti and the Dominican Republic. 
U.S. Nat. Mus. Bull: 155. 


Address: P. William Smith, P.O. Box 901341, Homestead, Florida 33090, U.S.A. 


© British Ornithologists’ Club 1997 


A.C. Vallely & T. Aversa 272 Bull. B.O.C. 1997 117(4) 


New and noteworthy bird records from Belize 
including the first record of Chestnut-collared 
Swift Cypseloides rutilus 


by Andrew C. Vallely & Thomas Aversa 
Received 18 October 1996 


Over thirty years have passed since Russell (1964) summarized what 
was then known about the status and distribution of birds in the 
Central American country of Belize. Among the more recent 
publications that have added to our understanding of avian dis- 
tributions in that area are those of Howell et al. (1992), Barlow et al. 
(1969, 1970, 1972), Mills & Rogers (1988), and Miller & Miller (1992). 

Since 1992, we have spent a combined total of 45 weeks in Belize. 
The following notes are all based on sight records made during the 
period January 1992 to April 1996. Included are the first record of 
Chestnut-collared Swift Cypseloides rutilus for Belize as well as notes on 
seven other species of poorly known or marginal occurrence in that 
country. It is noteworthy that while most of our time was spent in 
northern Belize, four of the eight observations detailed below resulted 
from less than four weeks spent in the poorly known southern half of 
the country. Further investigations in southern Belize will doubtless 
bring to light additional unusual records and range extensions (S. N. G. 
Howell pers. comm.). 


CINNAMON TEAL Anas cyanoptera 


Although described by Wood et al. (1986) as a very rare transient, 
Belize was not included in the range of this species by Howell & Webb 
Go): An adult male Cinnamon ‘Teal was observed feeding and 
roosting in flooded rice fields in the company of some 400 Blue- winged 
Teal A. discors and eight American Wigeon A. americana, approxi- 
mately 3km west of San Felipe, Orange Walk District, on 2 March 
1996. This bird was identified by its overall bright cinnamon coloration 
and red eye. 


BICOLORED HAWK Accipiter bicolor 


Russell (1964) listed three records of this species from Belize. Wood 
et al. (1986) described it as a very rare permanent resident in Belize. On 
9 April 1996, an adult Bicolored Hawk was observed carrying nesting 
material into a dense vine tangle along a logging road at Gran del Oro, 
Cayo District. As far as we know, this observation represents the first 
evidence of nesting by this little known species in Belize. 


CHESTNUT-COLLARED SWIFT Cypseloides rutilus 


At least four adult Chestnut-collared Swifts were present in a 
concentration of approximately 40 Chaetura swifts (probably Chaetura 
vauxt) at The Cockscomb Basin Wildlife Sanctuary, Stan Creek 


A.C. Vallely & T. Aversa 273 Bull. B.O.C. 1997 117(4) 


District, on 15 March 1993. Identification of these birds was based on 
their clearly visible chestnut collars. It is possible that several other 
(uniformly dark) swifts, seen with these birds, may have been C. rutilus 
immatures. Subsequently, S. N. G. Howell (zm litt.) observed a single 
individual of this species on 22 March 1996 at Monkey Bay Wildlife 
Sanctuary, Belize District. Howell & Webb (1995) noted that C. rutilus 
seems to withdraw from montane areas during the northern winter and 
that there is some evidence for migratory behaviour in this species. As 
far as we know, these observations represent the first records of this 
species from Belize. 


BLUE-THROATED GOLDENTAIL Hylocharis eliciae 

The first record for Belize was reported recently from Cayo District 
(Mills & Rogers 1988). A single male of this species was closely 
observed on 3 April 1994 as it perched and called in a Heliconia thicket 
along a disturbed creek-bed near Columbia Village, Toledo District. 


WHITE-WINGED BECARD Pachyramphus polychopterus 

Wood et al. (1986) described this species as a rare permanent resident 
in the southern half of the country, and Howell et al. (1992) reported a 
single mist-netted bird from Gallon Jug. On 30 March 1996, a single 
male White-winged Becard was found in the company of a mixed flock 
at the edge of a pasture at Bermudian Landing, Belize District. 
Identification was based on the absence of a whitish supraloral mark, 
thus ruling out P. major. 


NASHVILLE WARBLER Vermivora ruficapilla 

Russell (1964) listed two sight records of this species (both perhaps 
involving the same individual) three days apart at Gallon Jug, Orange 
Walk District. Lacking a specimen, he considered the species 
hypothetical in Belize. Wood et al. (1986) described it as a “‘very rare”’ 
transient. IT’. Aversa observed a single individual feeding with a flock of 
migrants at the edge of a clearing at Bermudian Landing, Belize 
District, on 31 March 1996. The identification of this bird was based 
on its small size, all-yellow underparts, grey cap and whitish eye-ring. 


CERULEAN WARBLER Dendroica cerulea 


Russell (1964) listed three records of this species from Belize. Wood 
et al. (1986) considered it a rare transient and (incorrectly) winter 
resident. Parker (1994) hypothesized that the Maya Mountains may 
form an important staging area for this species during spring migration. 
We noted a single male feeding in the mid-story along a logging road 
during a brief visit to New Maria Camp, Cayo District, on 7 April 
1996. This date is consistent with Parker’s (1994) observations made 
during 3-13 April 1992 at Toledo District. 


BLUE SEEDEATER A maurospiza concolor 

Howell et al. (1992) listed several records of this species from 
Monkey Bay and reported a mist-netted bird from Bermudian 
Landing, Belize District. T. Aversa saw at least two individuals (at least 


A.C. Vallely & T. Aversa 274 Bull. B.O.C. 1997 117(4) . 


one male and one female) in bamboo thickets along the Belize River at 
Bermudian Landing on 30 March 1996. 


GRASSLAND YELLOW-FINCH Sicalis luteola 


Wood et al. (1986) described it as a rare permanent resident in Belize. 
Howell & Webb (1995) noted that this species is ‘“somewhat nomadic’’. 
We observed a flock of 40-50 Grassland Yellow-finches as they flew 
and called over rice fields 3km west of San Felipe, Orange Walk 
District, on 17 January 1994. Numerous subsequent visits to this site 
failed to produce further observations. 


Acknowledgements 


The authors thank D. Dyer, V. Giles, M. Kasprzyk and D. Nyzio for their 
companionship in the field. A. Vallely is grateful to Manomet Observatory For 
Conservation Sciences and Progam For Belize for making his fieldwork possible. P. 
Sweet and S. N. G. Howell commented on a draft of this paper. 


References: 

Barlow, J. C., Dick, J. A., Baldwin, D. H. & Davis, R. A. 1969. New records of birds 
from British Honduras. [bis 111: 399-402. 

Barlow, J. C., Dick, J. A. & Pendergast, E. 1970. Additional records of birds from British 
Honduras (Belize). Condor 72: 371-372. 

Barlow, J. C., Dick, J. A., Weyer, D. & Young, W. F. 1972. New records of birds from 
British Honduras (Belize), including a Skua. Condor 74: 486-487. 

Howell, S. N. G. & Webb, S. 1995. A Guide to the Birds of Mexico and Northern Central 
America. Oxford Univ. Press. 

Howell, S. N. G., Dowell, B. A., James, D. A., Behrstock, R. A. & Robbins, C. S. 1992. 
New and noteworthy bird records from Belize. Bull. Brit. Orn. Cl. 112: 235-244. 

Miller, B. W. & Miller, C. M. 1992. Distributional notes and new species records for 
birds in Belize. Occasional Papers of the Belize Natural History Society 1: 6—25. 

Mills, E. D. & Rogers, D. T. Jr. 1988. First record of the Blue-throated Goldentail 
(Hylocharis eliciae) in Belize. Wilson Bull. 100: 510. 

Parker, T. A. 1994. Habitat, behavior, and spring migration of Cerulean Warbler in 
Belize. American Birds, spring 1994. 

Russell, S. M. 1964. A distributional study of the birds of British Honduras. Orn. 
Monogr. no. 1, American Ornithologists’ Union. 

Wood, D. S., Leberman, R. C. & Weyer, D. 1986. Checklist of the birds of Belize. 
Carnegie Mus. Nat. Hist. Special Publ. 12. 


Address: Andrew C. Vallely, 132 High Avenue, Apartment 6, Nyak, NY 06269, U.S.A. 
Thomas Aversa, 1907 North West Sloop Place, Seattle, WA 98117, U.S.A. 


© British Ornithologists’ Club (1997) 


F. C. Lowen et al. 275 Bull. B.O.C. 1997 117(4) 


New and noteworthy observations on the 
Paraguayan avifauna 


by James C. Lowen, Robert P. Clay, Juan Mazar Barnett, 
Alberto Madrono N., Mark Pearman, 
Bernabé Lopez Lantus, Joe A. Tobias, Durwyn C. Liley, 
Thomas M. Brooks, Estela Z. Esquivel S Fane M. Reid 


Received 29 October 1996 


Paraguay is divided by the Rio Paraguay into two _ discrete 
biogeographical regions. To the west lies the Chaco, a vast expanse of 
thorn scrub forests and seasonally inundated palm savannas. To the 
east, the Oriente combines several distinct ecosystems, and its avifauna 
is correspondingly diverse (Hayes 1995). In the northeast lie cerrado 
(Brazilian cerraddo; defined by Stotz et al. 1996 as “open campo 
grasslands with a sparse cover of shrubs and trees’’), in the southwest 
lie natural grasslands and marshes, and to the east is the humid Interior 
Atlantic Forest. 

Habitat destruction, of both the humid subtropical forests and the 
virgin grasslands, along with disturbance through hunting, trapping, 
pollution and infrastructural development, is rapidly depleting 
Paraguay’s avifauna. Less than 13% of the country’s Atlantic Forest 
remains, and almost all of this is partially degraded (Bozzano & Weik 
1992, DOA 1996). The grasslands of central South America are 
severely threatened by conversion to agricultural uses and have been 
long neglected as a conservation issue (Collar et al. 1992). No less than 
24 of the bird species now recorded in Paraguay are listed as threatened 
with global extinction by Collar et al. (1994). 

From July to December 1995, we carried out over 2000 hours of 
ornithological fieldwork in a variety of habitats at fourteen sites in the 
Oriente and one site in the Chaco. The conservation results of this 
fieldwork have been published elsewhere (Lowen et al. 1996a, 1996b). 
From these surveys, and from other recent unpublished fieldwork, we 
present noteworthy observations of 43 species, about 7% of the 
country’s avifauna (645 species: Hayes 1995). 

We define “noteworthy”? as follows: (a) first reliable country 
records (seven species: Russet-crowned Crake Anurolimnas viridis, 
Ocellated Crake Micropygia schomburgkiu, Spot-tailed Nightjar 
Caprimulgus maculicaudus, White-winged Nightjar Caprimulgus 
candicans, Hellmayr’s Pipit Anthus hellmayri, Masked Tityra T7ityra 
semifasciata and Yellow-bellied Seedeater Sporophila nigricollis); (b) 
first reliable country records for 50 years (eight species); (c) species 
with five or fewer reliable country records (an additional 24 species); 
(d) first reliable records for the Oriente (two species, excluding new 
country records) or the Chaco (four species, excluding new country 
records). Hayes (1995) provides appropriate baseline data with which 
to categorise our records, and subsequent fieldwork (e.g. Madrono & 
Esquivel 1995, Madrono & Esquivel 1997) is cited where relevant. 


JF. C. Lowen et al. 276 Bull. B.O.C. 1997 117(4) 


We publish elsewhere (Lowen et al. in press) a reassessment of the 
abundance and distribution of birds in Paraguay, which includes first 
records of 26 species for five of the seven biogeographic regions into 
which Hayes (1995) divides Paraguay. 


Abbreviations, conventions and species accounts 

Dpto.= Departmento; MN=Monumento Natural; MNHNP=Museo 
Nacional de Historia Natural del Paraguay; PN=Parque Nacional; 
RNBM=Reserva Natural del Bosque Mbaracayu, Dpto. Canindeyt; 
RNP=Reserva Natural Privada; RVS=Reserva de Vida Silvestre. 

Where possible, records were formally documented in one or more of 
the accepted manners (e.g. good quality photograph or tape-recording) 
and this is indicated in the text. For all mist-netted birds, full biometric 
data and a blood sample were taken before being photographed and 
released. Photographs and biometrics are available from the authors. 
Blood samples have been deposited with the Centre for Tropical 
Biodiversity, Copenhagen. Tape-recordings have been deposited at the 
National Sound Archive, Wildlife Section, London. We distinguish 
single- and multi-person records, listing up to three observers in 
alphabetical order. Authors are indicated by their initials; other 
observers are named in full. We use “‘et al.’’ to indicate that more than 
three observers were involved. All sight/aural records are supported by 
extensive field notes. No specimens were taken. 

‘Taxonomy and nomenclature for non-passerines follows Clements 
(1991), for passerines Ridgely & Tudor (1989, 1994). Species order 
follows Clements (1991). In each account, we give the species’ global 
status (if listed in Collar et al. 1994) and its status in Paraguay 
(according to Hayes 1995). We follow this with our records and 
summarise their significance. In certain cases, we include a brief 
discussion of status or identification features. For all localities 
mentioned, Appendix 1 lists details of department and co-ordinates. All 
sites are at low altitude (100-450 m). 


First reliable records of seven species for Paraguay 
RUSSET-CROWNED CRAKE Anurolimnas viridis 


Occurs from the Guianas and Venezuela to Brazil, where it is found 
south to Rio de Janeiro and Mato Grosso states (Sick 1993). Hayes 
(1995) does not list A. viridis for Paraguay. At Aguara Nu in the 
RNBM, Dpto. Canindeyt,, two calling birds were seen and tape- 
recorded on 19-20 September 1995, and one on 21 September (K. 
Zyskowski et al.). What were probably the same two birds were 
recorded at the same site 6-13 December 1995 (BLL). Both individuals 
were in an area of scrubby secondary growth on a dry substrate, within 
20m of an extensive marsh adjacent to a river. Both responded to 
playback, calling repeatedly and occasionally emerging from cover. 
Anurolimnas viridis has recently been found in Bolivia (Bates et al. 
1992); its range may be expanding with the spread of its favoured 
secondary growth habitat. 


F.C. Lowen et al. 277 Bull. B.O.C. 1997 117(4) 


OCELLATED CRAKE Micropygia schomburgkiu 

This near-threatened grassland rallid has an extensive range in South 
America (Collar et al. 1994). In Brazil, it occurs locally in Goids, Mato 
Grosso, Bahia and Sao Paulo states (Sick 1993). In Bolivia, it has 
recently been found to be widespread in a variety of habitats (Parker 
et al. 1991, Pearce-Higgins et al. 1995). M. schomburgkiu was not 
previously known from Paraguay, although it has been erroneously 
listed for the country (Hayes 1995). 

We recorded the species at two sites in the RNBM, Dpto. 
Canindeyu. At Lagunita, a minimum of eight calling individuals were 
present 9-13 September 1995 (MP et al.). One individual was 
tape-recorded and lured into mist-nets with playback. A photograph 
appears in Lowen et al. (1996a). Birds had first been heard, but not 
identified, on 21 November 1994 (RPC, AMN). At Aguara Nu, at least 
two individuals were heard (and tape-recorded) 14-19 September 1995 
(JMB ez al.). All birds were in tall dry grassland (often adjacent to 
wetter areas or lakes) and dry campo limpio type cerrado. 'The nearest 
known population is in Mato Grosso state, Brazil (Sick 1993). 
Micropygia schomburgki is likely to be rare in Paraguay, but may well 
be found in similar habitat further north in Dptos. Amambay and 
Concepcion. 


SPOT-TAILED NIGHTJAR Caprimulgus maculicaudus 


Caprimulgus maculicaudus has a disjunct range from Mexico south to 
north Bolivia and southeast Brazil (Hilty & Brown 1986, Howell & 
Webb 1995). In Brazil it is widespread but local, with only a few 
records from Mato Grosso south to Espirito Santo, Rio de Janeiro, Sao 
Paulo and Parana (Sick 1993). Its presence in Sao Paulo has recently 
been reconfirmed (Willis & Oniki 1993). Hayes (1995) does not list it 
for Paraguay. _ 

At Aguara Nu in the RNBM, Dpto. Canindeyt, two birds were 
calling 14-18 September 1995 (AMWN et al.). A male was mist-netted on 
16 September. It was photographed in the field and then recaught by 
hand on 17 September. This photograph is published in Lowen ez al. 
(1996a). All records were in a wet marshy valley and on adjacent drier 
hill slopes adjacent to gallery forest. 


WHITE-WINGED NIGHTJAR Caprimulgus candicans 

This poorly known nightjar, considered Critically Endangered by 
Collar et al. (1994), is only known from two old specimens in the 
grasslands of Mato Grosso and Sao Paulo in central Brazil, a series of 
modern records from Emas National Park in Goias, Brazil, and a 
specimen from Yucuma province, Bolivia, in 1987 (Collar et al. 1992, 
Sick 1993, Davis & Flores 1994). Hayes (1995) lists it as “‘hypothetical’’ 
for Paraguay. In common with most authors (e.g. Collar et al. 1992), 
Hayes rejects the reports by Azara (1802-5), which lack details of 
localities and dates. 

At Aguara Nu in the RNBM, Dpto. Canindeyt, we found 
Caprimulgus candicans to be not uncommon in cerrado, with records 


14—20 September (JMB et al.) and 6-13 December 1995 (BLL). At 


J. C. Lowen et al. 278 Bull. B.O.C.. 1997 117(4) 


least seven different males were recorded in the first hour after dusk on 
20 September, and a minimum of 14 birds were found during 
December surveys. On 20 September and 6—13 December seven birds 
(of two plumage types) were caught by hand. Photographs of one 
September male are published in Lowen et al. (1996a) and Tobias & 
Lowen (1996). A male was photographed in the field on 19 and 20 
September. Males appear to produce mechanical noises during display 
and these were tape-recorded on 10 December. Birds have again been 
recorded at Aguara Nu during 1996, with the first observation on 
27 August (EZE). 

These are the first documented records for Paraguay. However, 
Azara’s (1802-5) published description is identical to our males, and we 
consider that his records should be treated as genuine, albeit without 
locality. ‘There may be as many as 20 males in the 8000 ha of suitable 
habitat at Aguara Nu (although only 5500 ha is protected), and the site 
is evidently of critical importance for the conservation of this species 
(Lowen et al. 1996b). Birds showed a predilection for recently burnt 
areas; their ecological requirements would repay careful study. 


MASKED TITYRA Tityra semifasciata 


Widespread, occurring from northern Mexico south to Bolivia and 
Amazonian Brazil (Ridgely & Tudor 1994). Hayes (1995) does not list 
T. semifasciata for Paraguay. 

The first record for Paraguay comes from the Carapa waterfall trail 
in the northeast of the RNBM, Dpto. Canindeyu, where six birds were 
watched closely on 6 August 1995 (D. Finch et al.). Birds were seen 
again in this area during 1996 (EZE, AMN). At PN Cerro Cora, Dpto. 
Amambay, a single male was seen well but briefly on 10 October 1995 
(JMB, EZE, BLL). The Cerro Cora bird was easily distinguished from 
the congeneric Black-crowned Tityra T. inquisitor and Black-tailed 
Tityra T. cayana by its strikingly whitish upperparts and crown, broad 
white terminal tail band, restricted black forehead and facial area, and 
extensive red bill and bare skin around eyes. Being widespread over 
much of South America, it is not surprising that T. semifasciata should 
occur in Paraguay. 


HELLMAYR’S PIPIT Anthus hellmayri 


Anthus hellmayri occupies a disjunct range in the Andes of south 
Peru, west Bolivia, west Argentina and south Chile, with a separate 
population in southeast Brazil, Uruguay and east Argentina (Ridgely & 
Tudor 1989). Hayes (1995) lists it as “‘hypothetical’? in Paraguay, 
treating the two reports by Podtiaguin (1944) from the Chaco as 
unreliable: they lack plumage description and date. 

At RVS Yabebyry, Dpto. Misiones, we found up to ten birds daily 
28-31 October 1995 and estimated that six pairs were present (MP 
et al.). Most records were of birds singing from song posts (e.g. fences), 
a habit noted by Ridgely & Tudor (1989). Two individuals were 
photographed and two singing birds tape-recorded. These are the first 
documented records for Paraguay. It is likely to breed at RVS 
Yabebyry, and may have been overlooked in previous fieldwork in the 
south of the country. 


JF. C. Lowen et al. 279 Bull. B.O.C. 1997 117(4) 


YELLOW-BELLIED SEEDEATER Sporophila nigricollis 

Sporophila nigricollis is widespread in the lowlands from Costa Rica 
to north and east Bolivia, southern Brazil and northeast Argentina, 
though it is apparently absent from much of central Amazonia (Ridgely 
& Tudor 1989). Hayes (1995) does not list S. migricollis for Paraguay. 

At Puente Remanso, Dpto. Presidente Hayes, two adult males were 
seen on 3 November 1995 at the start of the Ruta TransChaco (M. 
Robbins, K. Zyskowski). In the Bahia de Asuncion, Dpto. Central, at 
least one male and a probable female were seen on 3 December 1995 
(RPC, EZE). These birds were in an overgrown garden in an area 
of marshy pools and low shrubs. Sporophila nigricollis is common 
and widespread north of Paraguay, and it is surprising that it has not 
been previously reported from the country. Further fieldwork will 
undoubtedly produce additional records. 


Eight species not reliably recorded in Paraguay for at least 
50 years 


GREY-BELLIED GOSHAWK Accipiter poliogaster 


Near-threatened (Collar et al. 1994). There have been six records in 
Paraguay, but none since 1938 (F. E. Hayes in litt. 1996). At RNP 
Itabo, Dpto. Canindeyu, pairs were seen on 15 (JCL) and 22 (JMB) 
October 1995, with singles on 17 and 18 October (BLL). All sightings 
were within 2 km of each other and presumably refer to the same birds. 


TINY HAWK Accipiter superciliosus 


Hayes (1995) lists three reliable records for Paraguay. All come from 
the Alto Parana region, in 1897 and 1934 (two). Hayes considers a 
recent record from Lago Ypacarai, Dpto. Central/Cordillera (Gonzales 
Torres & Gonzales Romero 1985), to be doubtful. At RNP Ypeti, 
Dpto. Caazapa, a single male was seen on 6 February 1994 (AMN). 
At Jejui-mi, RNBM, Dpto. Canindeyu, a female was seen well on 
30 September 1995 (RPC). The very small size and comparatively short 
tail of this forest Accipiter are distinctive: both observers are familiar 
with the species. 


PURPLE-WINGED GROUND-DOVE Claravis godefrida 


Critically Endangered (Collar et al. 1994). There are two records in 
Paraguay, both of pairs in Dpto. Alto Parana in the early 1890s (Bertoni 
1901). ‘The specimens have been lost (Hayes 1995). At RNP Itabo, 
Dpto. Canindeyu, a female was seen briefly but well on 6 December 
1994 (RPC). The bird was distinguished from female Blue 
Ground-dove Claravis pretiosa by its slightly larger size, lack of rufous 
rump and tail, off-white sides and white tips to splayed tail, and two 
obvious violaceous-brown wing-bars with darker borders. Additionally, 
a Claravts sp., probably C. godefrida, was heard (but not tape-recorded) 
in flowering Chusquea bamboo some 5 km away two days later. No 
additional records have been obtained during the 800 hours of 
fieldwork now conducted at the site (Brooks et al. 1993, Lowen ez al. 
1996b). 


F.C. Lowen et al. 280 Bull. B.O.C. 1997 117(4) 


Given the lack of other records from eastern Paraguay (e.g. not being 
found during bamboo flowerings in 1977; R. S. Ridgely in litt. 1991), it 
seems likely that Paraguay has always been on the edge of the species’ 
range. This record may represent a wandering individual either 
displaced by the disruption of the bamboo flowering cycle in Brazil or 
from a remnant population in Misiones province, Argentina (Lowen 


et al. 1996b). 


LARGE-TAILED ANTSHRIKE Mackenziaena leachiu 

Hayes (1995) lists three records for Paraguay, all of lost specimens 
from the Alto Parana region prior to 1936. At PN Caaguazu, Dpto. 
Caazapa, two individuals were seen and heard daily in different 
areas 21-23 July 1995 (TMB, RPC). At PN San Rafael, Dptos. 
Itapua/Caazapa, a pair was seen and heard 28-29 July 1995 (RPC 
et al.), with an additional bird heard on 29 July (JMR). All records 
were in bamboo-dominated secondary scrub on the forest edge. In 
Misiones province, Argentina, it is fairly common, favouring degraded 
forest and open forest areas (per JMB and MP). It was thus to be 
expected in southeast Paraguay. 


SPECKLE-BREASTED ANTPITTA Hylopezus nattereri 


Hylopezus natterent has recently been re-allocated full specific status 
(Whitney et al. 1995). Hayes (1995) lists just one record for Paraguay, 
from the Alto Parana region about 1903 (Bertoni 1904). At PN 
Caaguazu, Dpto. Caazapa, up to four individuals were calling in three 
separate areas daily 18-24 July 1995 (JCL et al.). One bird was 
tape-recorded distantly. At PN San Rafael, Dptos. Itapta/Caazapa, at 
least six individuals were calling in four areas on five dates 26—31 July 
1995. One bird was mist-netted on 31 July and a photograph appears in 
Lowen et al. (1996a). At Estancia Kaa’gua Rory, Dpto. Caaguazu, one 
or two birds were calling on 30 August, 2 and 7 September 1995 (RPC 
et al.). ‘These latter records may lend credence to local Aché indian 
reports of the species in the RNBM, Dpto. Canindeyt (AMN). 

The species’ presence in southeast Paraguay was to be expected; it is 
locally fairly common in bamboo-dominated forest in the adjacent 
Misiones province, Argentina (Benstead et al. 1993, MP). Whitney 
et al. (1995) note that, in the southern part of its range (including 
Paraguay), H. nattereri occurs at less than 300 m elevation, much lower 
than in the north. This pattern is reflected in other species with a 
similar geographical distribution, some of which are discussed in this 
paper (e.g. Large-tailed Antshrike and Bertoni’s Antbird). 


LESSER ELAENIA Elaenia chiriquensts 

Hayes (1995) only lists just one confirmed record in Paraguay (a 
specimen from the Alto Parana region in 1909) plus a hypothetical 
record in 1930. At Aguara Nu, RNBM, Dpto. Canindeyid, it was fairly 
common, with up to 12 seen ‘daily in cerrado 14-19 September 1995 
(JAT et al.). One bird was tape-recorded and photographed in the field. 
At the nearby Estancia Jiménez, Dpto. Canindeyu, two calling birds 
were seen in a small area of cerrado on 27 September 1995 (MP). 


¥. C. Lowen et al. 281 Bull. B.O.C. 1997 117(4) 


E. chiriquensis is probably under-recorded in Paraguay: it is an 
abundant passage migrant in the cerrado of Mato Grosso state, Brazil, 
during August and September (Sick 1993). 


BLACK-MASKED FINCH Coryphaspiza melanotis 


Vulnerable (Collar et al. 1994). Hayes (1995) lists one confirmed and 
one possible record for Paraguay, both prior to 1930. At Aguara Nu, 
RNBM, Dpto. Canindeyut, up to nine birds were encountered on each 
of six dates 14-21 September 1995 (JAT ez al.). These included at least 
five territorial males, all in an area of cerrado comprising tussocky grass 
never taller than 50 cm and small areas of bare ground. Unidentified 
scattered bushes and dwarf yata’1 palms (Butia sp.) were used as song 
perches; birds were otherwise terrestrial (Lowen et al. 1996b). One 
singing male was photographed and two birds tape-recorded. Several 
males were still singing during 6-13 December 1995 (BLL). Single 
juveniles were seen in both September and December, indicating 
breeding at the site. 

As Sick (1993) suggests for Brazil, C. melanotis is probably 
overlooked in Paraguay. It is likely to occur in similar cerrado habitat in 
Dptos. Amambay and Concepcion. 


RED-RUMPED WARBLING-FINCH Poospiza lateralis 


Hayes (1995) lists one certain record in Paraguay in 1911, plus one 
undated lost specimen. At PN Caaguazu, a male was watched at close 
range in scrub on the forest edge on 20 July 1995 (N. I. Etcheverry 
et al.). Renewed reports of this species were to be expected in southeast 


Paraguay, as it is locally common in a variety of habitats in northeast 
Argentina (JMB, MP). 


Species with five or fewer reliable records in Paraguay 
LEAST BITTERN /[xobrychus exilis 


Hayes (1995) lists five records for Paraguay. In the Chaco, we 
recorded Ixobrychus exilis at RNP La Golondrina, Dpto. Presidente 
Hayes: a male was seen well on 6 November 1995 (JCL). The first 
record for the Oriente came from PN Ypacarai, Dpto. Cordillera, 
where one was seen on 10 December 1995 (RPC, EZE, K. Zyskowski). 


CINEREOUS HARRIER Circus cinereus 
Hayes (1995) lists three reliable records from the Oriente and one 
hypothetical report from the Chaco. At Base Aerea, MN Bosque de 


Arary, Dpto. Itaptia, a pair were watched for a prolonged period on 
20 December 1995 (RPC, BLL). 


SWAINSON’S HAWK Buteo swainsoni 


Hayes (1995) lists just three records for Paraguay. At RNP Itabd, 
Dpto. Canindeyu, a light-phase bird was seen on 8 December 1994 
(RPC). At RNP Ypeti, Dpto. Caazapa, two light-phase birds were seen 


F.C. Lowen et al. 282 Bull. B.O.C. 1997 117(4) 


on 5 December 1995 (RPC). Two of the previous records for Paraguay 
were also in December (Hayes et al. 1990). 


GREY-BREASTED CRAKE Laterallus exilis 

Hayes (1995) lists two records: one in the Chaco in 1979 and one 
in the Oriente in 1994. At MN Bosque de Arary, Dpto. Itapta, 
three birds were calling daily (and one tape-recorded) at Puerto Turi, 
3—5 November 1995 (MP et al.). These add credence to Hayes’ (1995) 
speculation that it may breed in Paraguay. 


SPOT-WINGED PIGEON Columba maculosa 


There are only three acceptable Paraguayan records, all from the 
Chaco (Hayes 1995). The species is considered “‘hypothetical’’ in the 
Oriente (Hayes 1995). At RVS Yabebyry, Dpto. Misiones, seven birds 
were seen on 28 October, and two birds on 29 October 1995 (MP et al.). 
All birds were well seen flying over grassland with islands of forest. 
These are the first records for the Oriente. 


RED-SPECTACLED AMAZON Amazona pretret 


Endangered (Collar et al. 1994). This Atlantic Forest endemic is 
virtually restricted to Araucaria angustifolia forest in Rio Grande do 
Sul state, Brazil, with a few recent records from Misiones province, 
Argentina (Collar et al. 1994). Hayes (1995) lists Amazona pretrei as 
‘hypothetical’? in Paraguay despite three reports which include a 
well-described bird at RNP Itab6, Dpto. Canindeyu, on 15 August 
1992 (Brooks et al. 1993, 1995). At this site, which is a stronghold for 
the threatened Vinaceous Amazon A. vinacea, a pair were seen at close 
range on 2 August 1995 (DL). They were flushed several times from 
the subcanopy of open marshy woodland, always landing in dense 
vegetation. When seen at close range in flight, both birds showed a 
large red patch on the carpal joint, extensive red on the head, a clean 
green breast, and yellower vent. This combination of characters 
appears to eliminate all other Amazona species. 

Hayes (1995) treats A. pretrei as “‘hypothetical’’ in Paraguay in view . 
of the lack of formal documentation for existing reports and the lack of 
records from the adjacent Brazilian state of Parana. However, 
continuing to consider the species “‘hypothetical’’ in Paraguay may not 
be helpful for its conservation. Amazona pretrei moves northwards 
from Rio Grande do Sul during the austral winter (Collar et al. 1992), 
breeds in lowland riverine forest and utilizes several non-araucaria food 
sources (Varty et al. 1994). It is thus not inconceivable that stragglers 
reach Paraguay (where Araucaria angustifolia is now all but absent) 
fairly regularly in winter. 


TAWNY-BROWED OWL Pulsatrix koentswaldiana 

There are three records for Paraguay (Hayes 1995). These comprise 
an undated, historical specimen (Bertoni 1901), now lost (Hayes 1995), 
and 1992 records at sites in Dptos. Canindeyi and Caazapa (Brooks 
et al. 1993). We recorded P. koeniswaldiana at one of the 1992 sites: 
RNP Itab6, Dpto. Canindeyt. Up to four individuals were heard and 


JF. C. Lowen et al. 283 Bull. B.O.C. 1997 117(4) 


seen on six dates 17—23 October 1995 (DL et al.). Calls of at least three 
individuals were tape-recorded. This species is apparently rare in 
Paraguay, and may be restricted to humid forests of the extreme east. 


BLACK-BANDED OWL Strix huhula 

Hayes (1995) treats S. huhula as ‘hypothetical’? in Paraguay, 
considering the 1992 aural records published by Brooks et al. 
(1993, 1995) for RNP Itabo, Dpto. Canindeyu, to be insufficiently 
documented. Ericson & Amarilla (1997) subsequently described the 
first confirmed record for Paraguay: a female collected on 30 September 
1993 at PN Caaguazu, Dpto. Caazapa. We have recorded S. huhula 
during subsequent fieldwork at RNP Itabo. On 20 August 1993, a bird 
was calling repeatedly, but not tape-recorded (AMN). In 1995, up to 
two pairs were seen and heard in separate areas 20—22 October (MP et 
al.). All four individuals were tape-recorded and one photographed. 
The species has now been recorded in four distinct areas at RNP Itabo; 
it is probably a rare resident of forests in easternmost Paraguay. 


RUSTY-BARRED OWL Strix hylophila 

There are only three acceptable (and two very questionable) records 
in Paraguay, all from the Oriente (Hayes 1995). We found S. hylophila 
at two sites. At PN Caaguazt, Dpto. Caazapa, up to three birds were 
calling on six dates 15-23 July 1995, and a single bird was watched 
at close range on 16 July (JCL et al.). At PN San Rafael, Dptos. 
Itapua/Caazapa, single birds were heard calling on 26 and 30 July 1995. 


SICKLE-WINGED NIGHTJAR Lleothreptus anomalus 

Near-threatened (Collar et al. 1994). Hayes (1995) lists two 
confirmed records for Paraguay and a further three unconfirmed 
reports. There is a previously unpublished record of a female 
collected at a now flooded area of Isla Yacyreta (MNHNP 000566) on 
14 March 1992 (L. Amarilla im litt. 1996). We recorded the poorly 
known Eleothreptus anomalus at RNP Sombrero, Dpto. Cordillera. On 
25 August 1995, a male was seen and photographed in the field, then 
trapped (JCL et al.); a photograph appears in Lowen et al. (1996a). It 
was found in tall, dry grassland within 500 m of an extensive seasonally 
inundated marsh, and within 2 km of dry, open woodland. Available 
evidence suggests that this species is a rare but regular inhabitant of 
marshy grasslands of eastern Paraguay. The paucity of reports 
presumably stems from a patchy distribution and the generally silent, 
inconspicuous nature of the species. 


BLACK JACOBIN Melanotrochilus fuscus 

Hayes (1995) lists two previous Paraguayan records in 1987 and 
1994. The species has been recorded subsequently at the 1994 site (N. 
Pérez verbally 1995). At PN Caaguazu, Dpto. Caazapa, one bird was 
watched at close range on 16 July 1995 (RPC). There are no confusion 
species for this distinctive hummingbird. 


RUFOUS-THROATED SAPPHIRE Hylocharis sapphirina 


Brooks et al. (1993, 1995) detail two single observer sight records 
from the Oriente in 1992, but Hayes (1995) preferred to list it as 


F. C. Lowen et al. 284 Bull. B.O.C. 1997 117(4) 


‘“‘*hypothetical’”’ in Paraguay. He considered that the ‘“‘species is almost 
certainly present ... [but] further documentation is needed before it 
can be accepted’’. We observed it at three localities during 1995 
fieldwork. At PN Caaguazu, Dpto. Canindeyu, a male was watched at 
close range on 16 July 1995 (R. Barnes). At RNP Sombrero, Dpto. 
Cordillera, a male and a probable female were seen well on 28 August 
1995 (JCL). At PN Cerro Cora, Dpto. Amambay, a male gave close 
views on 10 October 1995 (JMB, BLL). 

The following plumage characters were common to all males seen: 
bill predominantly dark, but with varying amount of red; throat and 
centre of breast iridescent pale greenish-blue, grading to iridescent 
green on remainder of underparts; upperparts slightly darker green; 
tail and upper tail coverts bronze, tinged rufous. The Sombrero 
and Caaguazui males appeared to have a darker blue throat and 
more iridescent chestnut tail. Female Hylocharis sp.—probably H. 
sapphirina—with a slightly forked bluish tail, obvious white tips to the 
rectrices and more extensive rufous chin, were seen at Sombrero (as 
above), in the RNBM at Jejui-mi on 7 November 1994 (AMN) and at 
RNP Itabo, Dpto. Canindeyut, on 7 December 1994 (RPC). Hylocharis 
sapphirina has now been recorded over a wide area of the Oriente, and 
it is unfortunate that no documentation has been obtained. 


PYGMY KINGFISHER Chloroceryle aenea 


Hayes (1995) lists only two records for Paraguay, both from the 
Chaco. There are three subsequent records from the Oriente: at the 
RNBM, Dpto. Canindeyt in 1994 and 1995 (Madrono & Esquivel 
1997). We recorded it at Estancia Jiménez, Dpto. Canindeyt. Two 
birds were seen on 28 and 30 September 1995, with a female 
mist-netted on 29 September (JCL et al.). This site lies 20 km 
downstream of the RNBM. All reports of C. aenea in Paraguay come 
from the north, but given the two recent records of it in Misiones 
province, Argentina (Castelino 1990, MP), it may also occur further 
south in Paraguay. 


CANEBRAKE GROUNDCREEPER Clibanornis dendrocolaptoides 

Near-threatened (Collar et al. 1994). There are two records for 
Paraguay (Hayes 1995), including one in 1992 (Brooks et al. 1993, 
1995). At PN Caaguazt, Dpto. Caazapa, a single bird was seen in 
streamside bamboo on 18 July 1995 (JCL). Despite mist-netting in 
suitable habitat at several sites, and extensive voice playback (especially 
at RNBM), there were no further records of this Atlantic Forest 
endemic. This suggests it is very rare and probably restricted to the 
Parana basin of southeast Paraguay. 


WHITE-BROWED FOLIAGE-GLEANER Anabacerthia amaurotis 
Near-threatened (Collar et al. 1994). This Atlantic Forest endemic 
occurs in montane southeast Brazil (Parker et al. 1996, Stattersfield 
et al. 1997) and Misiones province, Argentina, where it is rare, but 
probably under-recorded (JMB). Hayes (1995) considered it “hypo- 
thetical’’ in Paraguay, listing eight sight records from three localities 


F. C. Lowen et al. 285 Bull. B.O.C. 1997 117(4) 


since 1992 (e.g. Brooks et al. 1993), but believing that “further 
documentation is needed’’. An additional sight record from the 
RNBM, Dpto. Canindeyt, in June 1995 (Madrono & Esquivel 1995) is 
best considered hypothetical (AMN). 

At PN Caaguazi, Dpto. Caazapa, we had seven sight records in 
different areas. Single birds were seen on 16 (R. Barnes), 17 (JAT), 18 
(RPC), 20 (JAT), 21 (JCL, DL) and 23 July 1995 (R. Johnson), and 
two on 22 July (R. Barnes). At PN San Rafael, Dptos. Itapua/Caazapa, 
a single was seen on 26 July (JAT) and two on 30 July 1995 (RPC). 

We consider that the likelihood of confusion with Buft-browed 
Foliage-gleaner Syndactyla rufosuperciliata (e.g. Hayes 1995) has 
been exaggerated. Anabacerthia amaurotis is considerably bulkier, 
with a far broader and whiter supercilium contrasting with thick 
dark eyestripe and crown. Its bright rufous tail contrasts with the 
brownish upperparts, while the straight bill is pale with a dark 
culmen. Additionally its vocalisations and behaviour differ from S. 
rufosuperciliata. 

There is an evident pattern to the 1992/1995 records, all occurring in 
the Parana basin forest of southeast Paraguay. This pattern is reflected 
in other species of montane southeast Brazil that are now being 
(re-)discovered in the lowland forests of southeast Paraguay (e.g. 
Large-tailed Antshrike, Bertoni’s Antbird and Diademed Tanager: see 
also Whitney et al. 1995). We suggest that A. amaurotis is a rare 
resident of the remaining forests of southeast Paraguay. 


BLACK-BILLED SCYTHEBILL Campylorhamphus falcularius 


Hayes (1995) lists three records, all from the Oriente, in 1893 (two) 
and 1977. At PN San Rafael, Dptos. Itapta/Caazapa, six individuals 
were seen on 26 July 1995, singles on 27, 29 and 30 July, and a bird 
mist-netted on 29 July (RPC et al.). 


RUFOUS-CAPPED ANTSHRIKE Thamnophilus ruficapillus 


Hayes (1995) lists only one reliable record (and three unconfirmed 
reports) from the Oriente. We recorded Thamnophilus ruficapillus at 
two sites. At Estancia Kaa’gua Rory, Dpto. Caaguazu, a pair (with 
the male singing) was seen in scrub on 29 August 1995 (JAT). A 
Thamnophilus sp. at Aguara Nu, RNBM, Dpto. Canindeyu, on 16 and 
18 September 1995 was initially identified as T. ruficapillus (see Lowen 
et al. 1996b). However, this individual is best left unidentified, now 
being thought to be closest to Rufous-winged Antshrike T. torquatus; 
further details will be published in due course. The paucity of 
Paraguayan records of T. ruficapillus is inexplicable. In northern 
Argentina, it is common in a variety of habitats including secondary 
growth (MP). 


BERTONI’S ANTBIRD Drymophila rubricollis 

Hayes (1995) lists just one record from Paraguay (in 1894: Bertoni 
1901), but Ridgely (1991) has also observed it at RNP Itab6, Dpto. 
Canindeyi, in June 1991. At PN Caaguaza, Dpto. Caazapa, a bird was 
seen and heard on 19 July 1995, and two were heard on 20 July (JAT 


F.C. Lowen et al. 286 Bull. B.O.C. 1997 117(4) 


et al.). At PN San Rafael, Dptos. Itapua/Caazapa, a bird was heard on 
30 July 1995 (JAT). At RNP Itabo, a bird was heard on 3 August 1995 
(JAT). All records were in dense scrub and forest islands on the forest 
edge. Drymophila rubricollis is common in northern Misiones province, 
Argentina, where it favours dense understorey growth, usually with 
Chusquea bamboo (JMB, MP). It was to be expected in eastern 
Paraguay, but is still only known from the extreme east. Although all 
recent records have been in the austral winter, this probably reflects the 
timing of fieldwork, rather than a genuine absence during the austral 
summer. 


COLLARED CRESCENT-CHEST Melanopareia torquata 


Hayes (1995) lists just one record for Paraguay, a 1938 specimen 
from Dpto. Amambay. Madrono & Esquivel (1997) provide the first 
recent record, with a bird seen in February 1995 at Aguara Nu in 
the RNBM, Dpto. Canindeyu. We found it in dwarf palm (Butia 
sp.) cerrado at the same site 14-20 September 1995 (JAT e al.). At 
least six pairs were present, and observed daily. Four males were 
singing and at least one pair were holding territory in a small group 
of palms. The species was still present in 1996, with many birds 
singing on 15 September (AMN). Melanopareia torquata is 
presumably a breeding resident here, and is likely to be found in 
other areas of similar habitat further north in Paraguay. We noted 
much variation in the colour and extent of nape collars, with at least 
one bird apparently lacking any collar. Such variation has not 
been reported in the identification literature (e.g. Ridgely & ‘Tudor 
1989). 


OCHRE-BREASTED PIPIT Anthus nattereri 


Endangered (Collar et al. 1994). Hayes (1995) lists up to four records 
for Paraguay. There are two records from the Chaco, which Hayes 
considers to be considerably out of range. There are two records in the 
Oriente, the most recent in 1977 (Ridgely & Tudor 1989). We found 
A. nattereri common at RVS Yabebyry, Dpto. Misiones, with 14 
singing birds in well-defined territories 27 October to 1 November 
1995 (BLL et al.). At the nearby MN Bosque del Arary, Dpto. Itapua, 
we found A. nattereri in two areas. At Base Aerea, it was common, with 
up to ten singing birds 1-3 November 1995 (MP et al.). At least four 
birds were still singing, and several other birds seen, on 20 December 
1995 (RPC, BLL). At Puerto Turi, a singing bird was seen and heard 
on 5 November 1995 (BLL). 

Different pairs were photographed in the field at RVS Yabebyry on 
28 and 30 October, and an additional singing bird was trapped 
on 1 November. The songs of at least two different birds were 
tape-recorded at this site, and two singing birds were tape-recorded 
at Base Aerea, MN Bosque de Arary, on 20 December 1995. The 
large populations of this very rare pipit at both sites are of very 
high conservation significance. It is fortunate that both sites receive 
at least nominal protection, covering a combined, and _ near- 
contiguous protected grassland area of over 40,000 ha (Lowen et al. 
1996b). 


F. C. Lowen et al. 287 Bull. B.O.C. 1997 117(4) 


GREEN-CHINNED EUPHONIA Euphonia chalybea 


Near-threatened (Collar et al. 1994). Hayes (1995) lists five certain 
and two possible records in the Oriente; one of these possible records 
(at the RNBM, Dpto. Canindeyu, in September 1992) should be 
considered certain (Brooks et al. 1993). There exists an additional 
specimen record (MNHNP 000323), previously labelled as a Palm 
Tanager Thraupis palmarum, taken at Puerto Triunfo, Dpto. Itaptia, on 
6 November 1985 (RPC, JCL, JAT). 

At PN Caaguazu, Dpto. Caazapa, we recorded a singing immature 
male or female on 18 July 1995 (RPC), an adult male on 20 July (JCL) 
and a presumed pair on 24 July (RPC). At PN San Rafael, Dptos. 
Itapuia/Caazapa, E. chalybea was fairly common, with 28 records 26-31 
July 1995; up to eleven birds were recorded daily in seven distinct areas 
(JAT ez al.). Most records were of pairs or singing males; one pair 
was photographed. At RNP Itab6, Dpto. Canindeyu, two female/ 
immatures were seen on 2 August 1995 (RB). The species is evidently 
fairly widespread, but rare, in eastern Paraguay, appearing most 
frequent in the southeast. 


DIADEMED TANAGER Stephanophorus diadematus 


Hayes (1995) lists five records, the four known localities lying in 
extreme eastern Paraguay. The only recent record is from 1992 (Brooks 
et al. 1993). At PN Caaguazt, Dpto. Caazapa, up to eight individuals 
were seen daily 16-22 July 1995, mostly in mixed flocks on the forest 
edge (RPC et al.). At PN San Rafael, Dptos. Itaptia/Caazapa, we 
recorded one bird on 27 July 1995 (JAT), and four on 29 July (R. 
Barnes, TMB, JCL), one of which was photographed. All birds were in 
scrub on the forest edge. This appears to be another member of the 
group of species (including Large-tailed Antshrike and Bertoni’s 
Antbird) that inhabit montane forest in southeast Brazil, but occur in 
scrubbier low-altitude habitat at the edge of their range in southeast 
Paraguay (see also Whitney et al. 1995). 


TEMMINCK’S SEEDEATER Sporophila falcirostris 


This bamboo specialist is considered Endangered by Collar et al. 
(1994). Hayes (1995) lists just one Paraguayan record, in 1977. 
Madrono & Esquivel (1995) found three males at the RNBM, Dpto. 
Canindeyd, in June 1995. We recorded it in bamboo (Merostachys sp.) 
at PN Caaguazu, Dpto. Caazapa, where males were seen on 18 July 
(RPC) and 19 July 1995 (JCL). Both observers had recent experience of 
Sporophila falcirostris in southeast Brazil. This rare species probably 
survives in low numbers in eastern Paraguay, perhaps being locally 
nomadic and moving between areas of bamboo. None of the 1995 birds 
was associated with bamboo flowerings, but it is possible that birds 
were lingering following such an episode in 1994 (RPC). Given the 
widespread deforestation of the adjacent Brazilian states, it would seem 
likely that any populations of S. falcirostris in eastern Paraguay and 
northeast Argentina are isolated from southeast Brazil (Lowen et al. 


1996b). 


F. C. Lowen et al. 288 Bull. B.O.C. 1997 117(4) 


SOOTY GRASSQUIT Tiaris fuliginosa 

Hayes (1995) lists one record for Paraguay, a bird at the RNBM, 
Dpto. Canindeyu, in 1992 (Brooks et al. 1993). Madrono & Esquivel 
(1997) detail additional records for the RNBM in September and 
November 1994. At Estancia Kaa’gua Rory, Dpto. Caaguazu, a female 
was mist-netted on 31 August 1995 (DL et al.). We recorded Tiaris 
fuliginosa in two areas at the RNBM. At Jejui-mi, a male was seen on 
8 September 1995 (JCL). At Lagunita, two males were found on 11 
(RPC), one male on 12 (RJ, JR) and a pair on 13 September 1995 
(BLL, JMB). Prior to 1992, the nearest known populations to Paraguay 
were in central Mato Grosso and in eastern Sao Paulo states, Brazil 
(Ridgely & Tudor 1989, Sibley & Monroe 1990). Subsequently, 
however, it has also been found in Misiones province, Argentina 
(Mazar Barnett & Herrera 1996). It would seem likely that T. fuliginosa 
occurs in low numbers throughout eastern Paraguay and northeast 
Argentina, but has—until recently—been overlooked. 


First reliable records for the Oriente or Chaco regions 
LEAST BITTERN Ixobrychus exilis 
See above. 


SPOT-FLANKED GALLINULE Gallinula melanops 

‘There are many records from the Chaco, but it is considered 
‘hypothetical’? in the Oriente (Hayes 1995), with just one old 
unconfirmed report lacking site and date details (Podtiaguin 1945). We 
recorded Gallinula melanops on the main lagoon at Lagunita, RNBM, 
Dpto. Canindeyu, where one was seen and photographed 10-13 
September 1995 (JMB et al.). This is the first documented record for 
the Oriente. 


SPOT-WINGED PIGEON Columba maculosa 
See above. 


MARSH SEEDEATER Sporophila palustris 


Endangered (Collar et al. 1994). Hayes (1995) only lists it for the 
Oriente. At RNP Golondrina, Dpto. Presidente Hayes, an adult male 
was present with a male Rufous-rumped Seedeater S. hypochroma on 7 
November 1995 (JCL). This is the first record for the Chaco. The date 
is that of a migrant, and a Chaco record is not surprising. Sporophila 
palustris usually arrives on breeding grounds in Corrientes province, 
Argentina, in the first week of November (Pearman & Abadie in 
press). 


CHESTNUT SEEDEATER Sporophila cinnamomea 


Near-threatened (Collar et al. 1994). Hayes (1995) gives a total of five 
records for Paraguay; additionally, a bird was recorded at RNBM, 
Dpto. Canindeyt, in October 1994 (Madrono & Esquivel 1995). At 
RNP Golondrina, Dpto. Presidente Hayes, a single male was seen in a 


F.C. Lowen et al. 289 Bull. B.O.C. 1997 117(4) 


mixed Sporophila flock on 8 November 1995 (BLL). This is the first 
record for the Chaco, and presumably refers to a bird moving south to 
breed in northern Argentina. 


LESSER GRASS-FINCH Emberizoides ypiranganus 


Near-threatened (Collar et al. 1994). Hayes (1995) lists just six 
records for Paraguay, all from the Oriente. The species was fairly 
common at RNP Golondrina, Dpto. Presidente Hayes, with up to ten 
individuals daily 7-9 November 1995 in wet grassland. These are the 
first records for the Chaco. The species probably occurs in similar 
habitat at other sites in the area. With few previous records in 
Paraguay, EF. ypiranganus has now been found over a wide area of the 
country (Lowen et al. 1996b). 


Discussion 


In this paper we discuss the occurrence of 43 bird species which 
previous evidence indicated to be exceptionally scarce in, or absent 
from, Paraguay. 

Three species—Anurolimnas viridis, Caprimulgus —candicans, 
Caprimulgus maculicaudus—were unexpected additions to _ the 
Paraguayan avifauna. Their nearest known populations lie several 
hundred kilometres from Paraguay. Our records probably represent 
previously overlooked populations rather than recent range extensions. 
Future fieldwork may help to clarify the situation. 

Two other species new to Paraguay—Tityra semifasciata, Sporophila 
nigricolliix—are relatively common throughout their extensive distri- 
bution. Our records may represent recent range extensions or indicate 
that the species occur in very low numbers at the edge of their range. 
Alternatively they may simply reflect the paucity of fieldwork in the 
country. Further surveys will doubtless find both species, and the 
resulting pattern of records may enable adjudication. Other species 
could be extending their ranges into Paraguay, but the available data 
are too few to justify firm conclusions. Such range extensions could 
represent open country birds extending into newly deforested zones or 
birds displaced by habitat destruction in adjacent areas (as suggested 
for the recent arrival of Curl-crested Jay Cyanocorax cristatellus at 
RNBM: see Madrono & Esquivel 1997). 

Most other noteworthy records refer to species previously overlooked 
(due either to their secretive behaviour or perceived identification 
difficulties) or to species dependent on habitats with a limited 
distribution in Paraguay. Such habitats include cerrados, restricted to 
the extreme north of eastern Paraguay (with isolated patches, such as 
Aguara Nu, RNBM, extending further south), and the grasslands of 
southern Paraguay. Until our 1994 and 1995 fieldwork, there had been 
very few comprehensive ornithological surveys of these areas, with 
most records attributable to occasional fortuitous observations. Our 
records of species such as Micropygia schomburgkiu, Elaenia chiriquensis, 
Melanopareia torquata, Anthus hellmayri, Anthus nattereri and 


F. C. Lowen et al. 290 Bull. B.O.C. 1997 117(4) 


Coryphaspiza melanotis, which occur in similar habitat in adjacent 
Brazil and Argentina, were thus not unexpected. 

Similarly, although the humid forest originally covered 55% of the 
Oriente (Bozzano & Weik 1992), there is a subset of Atlantic Forest 
endemics whose distribution in Paraguay appears restricted to the 
southeast, adjacent to Misiones province, Argentina (Lowen et al. 
1996b). Such species include Mackenziaena leachiu, Clibanornis 
dendrocolaptoides, Anabacerthia amaurotis, Campylorhamphus 
falcularius and Stephanophorus diadematus. Since recent ornithological 
fieldwork in the Paraguayan Atlantic Forest has concentrated on forests 
further north and west (e.g. Brooks et al. 1993, Madrono & Esquivel 
1997, Hayes & Scharf 1995), it is unsurprising that there have been few 
or no recent records of these species until renewed fieldwork at sites 
within their Paraguayan range. 

A number of the species are secretive and unlikely to be recorded 
if their vocalizations are not known. These include crakes, owls, 
Caprimulgus maculicaudus, Mackenziaena leachiu, Drymophila rubricollis 
and Hylopezus natterertz. Inadequate identification literature (e.g. for 
Anthus spp., Euphonia chalybea and Emberizoides ypiranganus) has 
a similar effect. Further fieldwork in appropriate habitat will prob- 
ably reveal these species as more widespread than current records 
suggest. 

Several migratory species may have been under-recorded due to a 
lack of fieldwork at the appropriate season. Buteo swainsoni, a North 
American breeder that spends the austral summer in Argentina, is 
likely to pass through Paraguay in reasonable numbers each year. 
Likewise, migrant Sporophila species such as S. palustris and S. 
cinnamomea probably pass through the Paraguayan Chaco each year en 
route between Brazil/Bolivia and Argentina. A few records may refer to 
vagrant individuals. This is perhaps the case for Gallinula melanops at 
RNBM, the single record of Poospiza lateralis, and both the 1992 
(Brooks et al. 1995) and 1995 records of Amazona pretret. 

New distributional information was gathered for seven globally 
threatened species, including the Critically Endangered Claravis 
godefrida and Caprimulgus candicans (Collar et al. 1994). All are 
primarily threatened by habitat loss (Collar et al. 1992, 1994). They 
are—by definition—likely to be genuinely rare in Paraguay, although at 
sites which preserve pristine areas of their habitat they could be locally 
common (e.g. Anthus natterer1 at_two southern grassland sites and 
Coryphaspiza melanotis at Aguara Nu, RNBM). 

Most of our records reflect the paucity of knowledge about the 
distribution of birds in Paraguay rather than genuine rarity of the 
species concerned. However, given the continuing destruction and 
degradation of Paraguay’s natural resources, it is likely that many are 
declining. Recent fieldwork (e.g. Brooks et al. 1993, 1995, Lowen et al. 
1995, 1996a, 1996b, Madrono & Esquivel 1997) has highlighted the 
urgent need for more biological fieldwork. The rate at which 
Paraguay’s ecosystems are being destroyed will otherwise result in the 
loss of the country’s avifauna before much of it has even been 
documented. 


$F. C. Lowen et al. 291 Bull. B.O.C. 1997 117(4) 


Acknowledgements 


For their vital work and companionship in and out of the field, we thank Ignacio Avila, 
Roger Barnes, Vanessa Doria, Nubia Etcheverry, Rosalina Farina, Bolivar Garcete 
Barrett, Richard Johnson, Mirna Perrens, Ramon Villalba, Sergio Villanueva and Kristof 
Zyskowski. Luis Amarilla, Nubia Etcheverry, Davis Finch, Mark Robbins and Kristof 
Zyskowski kindly gave their permission to cite previously unpublished field observations 
and specimen records. 1995 fieldwork was conducted with the permission and help of the 
Fundacion Moisés Bertoni para la Conservacion de la Naturaleza, the Direccion de 
Parques Nacionales y Vida Silvestre, the Facultad de Ciencas Exactas y Naturales and the 
Museo Nacional de Historia Natural del Paraguay. Additional logistical support was 
provided by the British Embassy, Asuncion. Individuals from these institutions are 
credited in Lowen et al. (1996b). RPC’s field work in 1994 was funded by a British 
Ecological Society Research Travel Grant. Our 1995 fieldwork was part-funded by grants 
from the BP Conservation Expedition Competition, Whitley Animal Protection League, 
Peoples Trust for Endangered Species, Shell International, Royal Geographical Society, 
British Ornithologists’ Union, Bird Exploration Fund, British Ecological Society, 
Gilchrist Educational Trust, Harvard Travellers Club Permanent Fund and Institute of 
Biology. Other donors are credited in Lowen et al. (1996b). Nigel J. Collar and David W. 


Snow kindly reviewed the text, providing many useful suggestions. 


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importancia en la conservaciOn de aves amenazadas, cuasi-amenazadas y endémicas 
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Podtiaguin, B. 1945. Catalogo sistematico de las aves de Paraguay. Rev. Soc. Cient. 
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Ridgely, R. S. & Tudor, G. 1989. The Birds of South America: the oscine passerines. 
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Bull. B.O.C. 1997 117(4) 


Addresses: James C. Lowen, BirdLife International, Wellbrook Court, Girton Road, 


Cambridge CB3 ONA, U.K. Robert P. Clay and Joe A. Tobias, Dept. of Zoology, 
University of Cambridge, Downing Street, Cambridge CB2 3EJ, U.K. Juan Mazar 
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de la Naturaleza, Calle Procer Carlos Argiiello 208, CC714, Asuncion, Paraguay. 
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Biological Sciences, University of East Anglia, Norwich NR47TJ, U.K. Thomas 
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Knoxville, TN 37996-1610, U.S.A. Jane M. Reid, Newnham College, Cambridge, 
U.K. 


© British Ornithologists’ Club 1997 


Appendix 
A summary of the location of sites mentioned in the text 


Region Site Department Coordinates 
Oriente RVS Yabebyry Misiones 27°10'S, 57°00"W 
MN Bosque de Arary Itapua 27°27'S, 56°48°W 
/Base Aerea 20° 25'S 57 Vt AWwW 
/Puerto Turi 27°24'S, 57°14"'W 
RNP Ypeti Caazapa 25°33’'S, 55°30'W 
RNP Itabo Canindeyu 24°20'S, 54°35'W 
PN Caaguazu Caazapa 26°04'S, 55°45'W 
PN San Rafael Itapta 26°25’S, 55°40'W 
RNB Mbaracayu Canindeyu 24°07'S, 55°23"W 
/Jejui-mi 24°08’S, 55°31'W 
/Lagunita_ 24°08’S, 55°25'W 
/Aguara Nu 24°09’S, 55°16'W 
/Carapa waterfall 24°00’S, 55°20'W 
PN Ypacarai Central/Cordillera 25 15 9, 5,19 WW 
RNP Sombrero Cordillera 25°00'S, 56°38’W 
Estancia Kaa’gua Rory Caaguazu 24°46'S, 55°26’W 
Bahia de Asuncion Central 25°18'S, 57°40'W 
Estancia Jiménez Canindeyu 24°13’S, 55°38’W 
PN Cerro Cora Amambay 22°39'S, 56°00’W 
Chaco RNP Golondrina Presidente Hayes 24°59’S, 57°43'W 


Puerto Remanso 


Presidente Hayes 


25°12’S, 57°33’W 


M.A. Raposo & R. Parrini 294 Bull. B.O.C. 1997 117(4) 


On the validity of the Half-collared Sparrow 
Arremon semitorquatus Swainson, 1837 


by Marcos A. Raposo & Ricardo Parrim 
Received 9 November 1996 


Many authors have stressed that it is necessary to review the many 
existing polytypic species because of the conceptual problems 
presented by the subspecific rank (Rosen 1978, 1979, Nelson & Platnick 
1981, Wiley 1981, Cracraft 1983, 1992, McKitrick & Zink 1988). 

The Pectoral Sparrow Arremon taciturnus (Hermann, 1783) is a 
classical example of a problematic taxon. It is a Neotropical 
undergrowth forest dweller generally treated as consisting of four 
subspecies: A. t. nigrirostris Sclater, 1886, from Peru, Bolivia, and 
probably northwestern Argentina (Meyer de Schauensee 1982); A. tf. 
axillaris Sclater, 1854, from the eastern part of Colombia and adjacent 
Venezuela; A. t. taciturnus ranging from southern Venezuela, the 
Guyanas and the right bank of the rio Amazonas through the states of 
Mato Grosso, Goias, Tocantins, Para and southward to Minas Gerais 
and Espirito Santo; and A. t. semitorquatus Swainson, 1837 which is 
restricted to Brazil from Rio de Janeiro to Rio Grande do Sul 
(Hellmayr 1938, Pinto 1944). 

In spite of the well marked differences between the last two taxa their 
conspecificity was defended by Hellmayr (op. cit.) and Pinto (op. cit.) 
on the basis of a supposed intergradation zone, particularly at Serra 
dos Orgdos, a mountain range in Rio de Janeiro. ‘These authors 
distinguished A. t. semitorquatus from A. t. taciturnus by the colour of 
the lower mandible (yellow in the former and blackish-brown in the 
latter), the lesser upper wing coverts (nearly the same olive as the back 
in the former, bright yellow in the latter), and the black pectoral collar 
(interrupted at the centre of the neck in A. t. semitorquatus, complete in 
the nominate subspecies). The presence of an “‘intergradation zone’’ 
was based on the yariation of the two last characters in some specimens 
from Serra dos Orgdos and in one skin from. Ipanema (upland Sao 
Paulo). 

The purpose of this work is to re-examine the evidence adduced by 
Hellmayr and Pinto, and to review the taxonomic status of Swainson’s 
form. 


Methods 


We conducted an analysis of the specimens housed in the Museu 
Nacional of Rio de Janeiro and Museu de Zoologia da Universidade de 
Sao Paulo. A total of 243 skins were examined, 54 corresponding to the 
diagnosis of A. t. semitorquatus and 189 to that of the nominate form. 
The measurements taken were: bill length (calmen); wing length 
(chord); and tail length. Some field notes were made and specimens 


M.A. Raposo & R. Parrini 295 Bull. B.O.C. 1997 117(4) 


were collected in Bahia and Rio de Janeiro, mainly in Serra dos Orgios, 
the alleged intergradation zone between A. t. taciturnus and A. t. 
semitorquatus. The Colombian and Peruvian/Bolivian subspecies were 
not included in the present comparison since they are widely separated 
from A. t. semitorquatus, our main subject. A good diagnosis of those 
forms can be found in Hellmayr (1938). The geographical co-ordinates 
of all the localities mentioned here may be found in Paynter & Traylor 
(1991) and Vanzolini (1992). 


Results 


Our analysis produced no evidence to corroborate the conspecificity of 
semitorquatus and taciturnus, nor the existence of the intergradation 
zone mentioned by Hellmayr and Pinto. No individual of A. tf. 
taciturnus was found in Rio de Janeiro (the proposed hybridisation 
zone) where we only found typical semitorquatus, which was also 
recorded northward to northeastern Espirito Santo (Santa Teresa, 
19°55’S, 40°36’W and Jatiboca, 20°05’S, 40°55’W) and eastern Minas 
Gerais (Mariana, 19°30’S, 41°00’W). It is noteworthy that this 
northward extension was not mentioned by those authors, and that 
it makes the intergradation zone at the interior of Rio de Janeiro 
state incomprehensible (see Fig. 1). The apparent allopatry of these 
two forms and the clear distinction between them, although their 
ranges abut, indicates that semitorquatus should be elevated to species 
rank. 


Diagnosis of A. semitorquatus 

The characters presented by Hellmayr and Pinto for the differen- 
tiation of A. semitorquatus and A. taciturnus were found to be constant. 
A. semitorquatus is characterised by the presence of an interrupted black 
collar, in contrast to the complete one found in adult males of nominate 
A. taciturnus. In A. semitorquatus, the bill had a yellow mandible 
contrasting with a blackish-brown maxilla, and the lesser upper wing 
coverts are nearly of the same colour as the rest of the dorsal plumage, 
while A. taciturnus has an uniform blackish bill and yolk-yellow lesser 
upper wing coverts. The sides of the belly in semitorquatus are invaded 
by dark grey, usually not present in A. taciturnus. The females of 
semitorquatus have an interrupted collar like that of the males, contrary 
to the accentuated dimorphism presented by A. taciturnus, the females 
of which do not have the black collar or have this feature reduced to a 
slight and inconspicuous grey jugular band. In both species, the 
females generally have a clear suffusion of buff on the ventral area, most 
pronounced on the throat. 

The other species that could be mistaken for A. semitorquatus is 
Arremon flavirostris Swainson, 1837, notably the nominate subspecies 
which, like the former, has bright olive upper parts. The latter species 
has an orange bill with a black ridge, a white chin and a complete black 
pectoral band in both sexes. A. semitorquatus and A. taciturnus both 
have a black chin-spot and a white superciliary stripe reaching to the 
base of bill, while A. flavirostris and its sister-taxon A. polionotus 


M.A. Raposo & R. Parrin 296 Bull. B.O.C. 1997 117(4) 


Figure 1. Map showing the distributions of A. semitorquatus (O) and A. taciturnus (@) in 
eastern South America after our studies, and the proposed intergradation area of 
Hellmayr (1938). 


Bonaparte, 1851, both occurring in central Brazil, have the white 
superciliary stripe starting just above the eye (Silva 1991). A. polionotus 
has, moreover, dark grey upper parts. A. flavirostris dorbignii Sclater, 


M.A. Raposo & R. Parrini 297 Bull. B.O.C. 1997 117(4) 


1856 (Bolivia to Argentina) resembles the nominate subspecies. The 
status of the Brazilian subspecies of the A. flavirostris complex was 
recently reviewed by Silva (op. czt.). 


Individual variation j 

One male (MN 38305) from Serra dos Orgaos (Nova Friburgo) 
shows an ill-defined collar similar to that described by Hellmayr for an 
intermediate between the interrupted and the complete collar patterns 
of semitorquatus and taciturnus. In spite of it, this specimen retains all 
the other characters of A. semitorquatus. The fact that it comes from the 
centre of the range of A. semitorquatus, far from that of A. taciturnus, 
leads us to suppose that this plumage pattern may be of the ontogenetic 
origin or represent a rare individual variation. This feature alone 1s not 
enough to justify the suggestion of intergradation. ‘Iwo specimens of 
A. taciturnus from Para (rio Cururu, 07°12'S, 58°03’W) show a similar 
variation, having an incomplete pectoral band. We have come across 
other cases of isolated individual variation. 


Measurements 

The examination of skins of both sexes of A. semitorquatus and A. 
taciturnus revealed some notable differences in their measurements. 
The tail of semitorquatus (mean 66.51; s.d. 2.81) is noticeably longer 
(t=12.73, P<0.01) than that of taciturnus (mean 58.33; s.d. 3.67). The 
wing chord of semitorquatus (mean 69.8; s.d. 3.23), in contrast to the 
tail, is usually shorter (t=3.27, P<0.01) than that of taciturnus (mean 
71.95; s.d. 3.49). The bill of semztorquatus (mean 12.32; s.d. 0.61) is in 
general shorter (t=6.36, P<Q.01) than that of taciturnus (mean 13.06; 
s.d. 0.51). 


Geographic distribution 

Based on the examined material, A. semitorquatus occurs from 
Espirito Santo (Santa Teresa and Jatiboca) and Minas Gerais 
(““Fazenda Taveira’ and Caparao), southward through Rio de Janeiro 
(Trés Rios, Cantagalo, Nova Friburgo, Teresopolis and Mangaratiba), 
Sao Paulo (Ribeirdo Fundo, Serra da Cantareira, Mogi das Cruzes, 
Cabreuva, Piquete, Rio Grande, Barra do Ribeiréo Onca Parda, 
Salesopolis and Itapetininga), Parana (Morretes), probably Santa 
Catarina, to Rio Grande do Sul (Novo Hamburgo). Rio de Janeiro was 
suggested as the type locality by Berlepsch (in Hellmayr 1938). 

In the states of Minas Gerais (Machacalis and Baixo Suassui) and 
Espirito Santo (e.g. Soretama, Ibiragu, Baixo Guandu and Rio Sao 
José) A. taciturnus occupies mainly lowlands, while A. semitorquatus 
seems to be confined to the mountains (e.g. Santa Teresa). This rule is 
not valid from Rio de Janeiro southward, where A. taciturnus is absent 
and A. semitorquatus also occurs in lowland areas (e.g. Mangaratiba and 
Morretes). The same phenomenon occurs with A. taciturnus, as in the 
northern portion of its range it is found in the mountains (Serra de 
Baturité, Serra da Neblina, etc.) as well as in the lowlands (e.g. Porto 
Seguro). 


M.A. Raposo & R. Parrini 298 Bull. B.O.C. 1997 117(4) 


Final remarks 


To conclude, Arremon semitorquatus is a good species, found to our 
knowledge only in Brazil. There is no geographic area where it has 
been found in sympatry with A. taciturnus. ‘The intergradation 
zone proposed by Hellmayr and Pinto may have been based on 
misinterpretation of individual variation in the size of the pectoral 
band. Phylogenetic relationships within this genus are still unclear, and 
a revision is desirable. ‘The presence of conspicuous sexual dimorphism 
in A. taciturnus (female without pectoral band) is an interesting feature 
that is not shared by any of its congeners. The taxonomic status of the 
subspecies of A. taciturnus should be scrutinised, especially A. t. 
nigrirostris which lacks good diagnostic characters. 


Acknowledgements 


We would like to thank Dante M. Teixeira, Jorge Nacinovic and the rest of the staff 
members of Setor de Ornitologia, Museu Nacional/UFRJ for reviewing and commenting 
on our manuscript. We are also grateful to CAPES for financial support and to Helio 
F. A. Camargo (MZUSP) for providing us with all facilities to study the series at Sao 
Paulo. 


References: 

Cracraft, J. 1983. Species concepts and speciation analysis. Current Ornithology 1: 
159-187. 

Cracraft, J. 1992. The species of the birds-of-paradise (Paradisaeidae): applying the 
phylogenetic species concept to a complex pattern of diversification. Cladistics 8: 
1-43. 

Hellmayr, C. H. 1983. Catalogue of birds of the Americas and the adjacent islands. Part 
XI. Publ. Field Mus. Nat. Hist., Zool. Ser. 13. 

McKitrick, M. & Zink, R. 1988. Species concepts in ornithology. Condor 90: 1-14. 

Meyer de Schauensee, R. 1982. A Guide to the Birds of South America. Livingston. 

Nelson, G. J. & Platnick, N. I. 1981. Systematics and Biogeography: Cladistics and 
Vicariance. Columbia Univ. Press. 

Paynter, R. A. & Traylor, M. A. 1991. Ornithological Gazetteers of Brazil (2 vol.). 
Harvard Univ. Press. 

Pinto, O. M. O. 1994. Catdlogo das Aves do Brasil e Lista dos Exemplares. Existentes na 
Colecao do Departamento de Zoologia. Pt. 2. Sao Paulo. 

Rosen, D. E. 1978. Vicariant patterns and historical explanation in biogeography. Syst. 
Zool. 27: 159-188. 

Rosen, D. E. 1979. Fishes from the uplands and intermontane basins of Guatemala: 
revisionary studies and comparative geography. Bull. Am. Mus. Nat. Hist. 162: 
267-376. 

Sick, H. 1985. Ornitologia brasileira: uma introdugdo. Ed. Univ. Brasilia. 

Silva, J. M. C. da. 1991. Geographical variation in the Saffron-billed Sparrow Arremon 
flavirostris. Bull. Brit. Orn. Cl. 111: 152-155. 

Vanzolini, P. E. 1992. A Supplement to the Ornithological Gazetteer of Brazil. 
Universidade de Sao Paulo, Sao Paulo. 


Wiley, E. O. 1981. Phylogenetics. The theory and practice of phylogenetic systematics. 
J. Wiley & Sons, New York. 


Address: Marcos A. Raposo, Setor de Ornitologia, Museu Nacional, Quinta da Boa Vista, 
Rio de Janeiro (RJ), CEP 20940-040, Brazil. 


© British Ornithologists’ Club 1997 


P. J. Cowan 299 Bull. B.O.C. 1997 117(4) 


What is a desert bird? 
by PF. Cowan 


Received 29 November 1996 


Desert is arid (low precipitation, high potential evaporation, rare and 
ephemeral liquid water accumulation on surface) and has a relatively 
small amount of vegetation. A bird in a desert might be resident, 
wintering, on migration or have some other temporal status. 
Spatially, some resident species appear characteristic of the desert 
whilst other avian desert residents have widespread distributions 
outside deserts. Some bird species occur in the desert proper whilst 
others are found in oases. 

Many studies of “‘desert birds’’ have been reported in the scientific 
literature, including recent reviews by Webster (1991) and Maclean 
(1996) on ecophysiological aspects of avian survival in deserts. 
However, defining a desert bird is problematic (Cowan 1990, Wiens 
1991) and the particular usage in a study is often not explicit. Lowe 
(1968) noted that animals referred to as desert species have often 
included species with a far wider distribution outside deserts. Lowe 
(1968) distinguished between non-desert species of animals, which 
occur in ‘“‘permissive’’ desert environments and are more wide- 
spread outside deserts, and desert species which are either ‘“‘desert- 
only” arid habitat species or “‘desert-included”’ species occurring in 
both desert-arid habitats and semi-arid regions outside deserts. The 
present note is intended to further elucidate the question: what is a 
desert bird? 

A desert bird could be a bird species that has evolutionary 
adaptations to the arid habitats of deserts (Maclean 1976), though any 
suggested adaptation needs to be proved. Bradshaw (1988) considered 
that desert reptiles survive due to general reptilian adaptations rather 
than the possession of unique adaptations to deserts. Birds may only 
need adaptive refinements to ensure survival in deserts due to 
favourable general avian adaptations (Miller 1963). ‘Thomas (1984a,b) 
suggested that desert birds have a multiplicity of relatively minor 
evolutionary adaptations to deserts the cumulative effect of which is 
substantial. The discovery that cutaneous evaporative cooling 
constitutes a major heat defence mechanism in certain columbid birds 
contrary to previous expectations (Webster 1991), suggests that 
significant individual adaptations of birds to the arid habitats of 
deserts may remain to be discovered. 

Adaptations to deserts need not be confined to species restricted in 
distribution to deserts, e.g. birds which migrate through desert regions 
might acquire desert adaptations (Miller 1963); but species restricted to 
the arid habitats of deserts presumably would be the most likely to 
possess significant desert adaptations. An ecogeographic definition of a 
desert bird therefore should aid in the selection of species for study in 
the hunt for avian adaptations to deserts. 


P. F. Cowan 300 Bull. B.O.C. 1997 117(4) 


What is a desert? 


The question of what is a desert bird includes the perennial problem of 
precisely defining what is a desert. Differing systems of discrete major 
ecosystem types have been used to describe and map the Earth’s 
terrestrial regions; tundra, desert, tropical rain forest and other major 
ecosystem types being variously defined using mainly climatic and 
vegetational criteria (e.g. Bailey 1983, 1989). Numerous other studies 
have concentrated on arid environments and these have used a wide 
variety of desert definitions involving, for example, vegetational 
criteria, the simple specification of isohyets and the use of aridity 
indices incorporating precipitation, temperature and other climatic 
variables (review by McGinnies et al. 1968). Some authors, e.g. Le 
Houérou (1977), have restricted their definition of a desert to include 
only the more extreme arid conditions. Schmidt (1979) presented 12 
different delineations of the Chihuahuan desert of North America and a 
13th, his version. The ecological zones of North Africa, from the 
central Sahara desert north to the 400 mm isohyet, presented by Le 
Houérou (1979) and based on climate, vegetation and land use, are, 
although he stated otherwise, arbitrary. Different, but equally valid, 
subdivisions of this region are possible (e.g. Ayyad & Ghabbour 1986). 
Only arbitrary criteria are available for the distinction between desert 
and the less arid major ecosystem types (e.g. Whittaker 1975, Beard 
1980, Monod 1986, World Conservation Monitoring Centre 1992). 
The desert concept is subjective and, therefore, any definition of a 
desert bird is also subjective. 

The UNESCO (1979) map of the world distribution of arid and 
semi-arid regions, based on aridity index values for meteorological 
stations but supplemented by taking soil, relief and vegetation data into 
account, is arguably the best general-purpose aridity map yet produced 
(Verstraete & Schwartz 1991). The aridity index of the UNESCO 
(1979) map is the ratio P/ETP, where P is the mean annual 
precipitation and ETP is the mean annual potential evapotranspiration 
calculated using the Penman formula. P/ETP less than 0.20 is arid. The 
arid (arid and hyper-arid) regions of this map, but not its semi-arid and 
sub-humid regions, can be considered to be deserts (Cowan 1987, 1990, 
1996, Oberlander 1994). 

The world’s deserts, using the UNESCO (1979) map, lie between 
latitudes 50°N and 50°S. Major desert areas occur in North and South 
America, northern and southern Africa, Asia, and Australia. Polar 
‘““deserts’” (Logan 1968) are excluded. The high altitude deserts of the 
Tibetan Plateau are included. The hyper-arid regions of the UNESCO 
(1979) map are mainly restricted to North Africa and Arabia and are 
absent from Australia. The arid (arid and extremely arid) areas of its 
predecessor map, produced by Peveril Meigs for UNESCO, were used 
by McGinnies et al. (1968) and Evenari (1985) for desert delimitation. 

Within the perimeter of a desert a distinction exists between desert 
and oasis. Oases include traditional oases, irrigated agricultural 
projects, cities and habitats associated with perennial rivers. Lakes 
or pools in the desert, with associated vegetation, can attract a 


P. ¥. Cowan 301 Bull. B.O.C. 1997 117(4) 


considerable avifauna of wetland species (e.g. Cowan 1985). Non-arid 
highlands can also occur within the desert perimeter (e.g. Schmidt 
1989). The arid habitats of deserts include the normally-dry valleys, 
which experience stream flow only after sufficiently heavy rain. Some — 
areas in oases, away from irrigation and water, appear effectively to be 
arid habitats. Also, previously irrigated agricultural areas return to 
desert-like conditions. 


Desert birds 


The ideal ecogeographic definition of a desert bird would be a species 
completely restricted in its breeding and non-breeding range to the arid 
habitats of deserts. Unfortunately, comparison of the distributions of 
avian species with the boundaries of the world’s deserts, the arid and 
hyper-arid regions of the UNESCO (1979) map, soon reveals a 
problem. Desert Lark Ammomanes deserti, Desert Wheatear Oenanthe 
deserti, ‘Trumpeter Finch Bucanetes githagineus and many others of 
those avian species which appear good candidates for meeting the 
requirements of the ideal definition (e.g. Casselton 1984), are not 
completely restricted to the arid habitats of deserts; they can also 
occur and breed in semi-arid regions outside the desert such as 
steppe, drier savanna or semi-arid mountains. Apparent desert birds 
are sometimes found in oasis situations; e.g. Spotted, Crowned and 
Lichtenstein’s Sandgrouse Pterocles senegallus, P. coronatus, P. 
lichtensteinit drink at permanent pools and village irrigation channels 
(Thomas & Robin 1977), Black-crowned Finch Larks Evemopterix 
nigriceps are abundant in late summer at a lush desert golf course 
(Richardson 1990), and in central Saudi Arabia, in late summer, local 
concentrations of Brown-necked Ravens Corvus ruficollis occur near 
water, including up to 500 at a reservoir on the outskirts of Riyadh 
(Jennings 1980). Some likely desert birds have occurred as vagrants 
ia, eOutmies (a considerable , distance” from” deserts, . e.g: 33 
Cream-coloured Coursers Cursorius cursor have been recorded in the 
British Isles, nine Desert Warblers Sylvia nana in Sweden 
(Lewington et al. 1991). Ignoring vagrancy, the Bar-tailed Desert 
Lark Ammomanes cincturus is probably the closest to meeting the 
ideal definition of the 25 species listed by Casselton (1984) as the 
desert-associated members of the Sahara desert avifauna. 

More realistically, desert birds can be considered as those species 
which occur primarily within the arid habitats of deserts, any 
occurrence elsewhere being considerably less important. This 
definition includes species which are completely restricted to the arid 
habitats of deserts but excludes, for example, species which occur 
equally in deserts and semi-arid regions. A desert bird could breed in 
one desert, migrate with stop-overs in less-arid regions, then 
overwinter in another desert. The imprecision of the terms “‘primarily”’ 
and “‘considerably”’ in the definition adds further subjectivity to any list 
of desert birds produced. Selection of species for a list of the desert 
birds of a desert involves assessing from the literature the world 
distribution of each of the avian species recorded there. I listed 23 (later 


P. F. Cowan 302 Bull. B.O.C. 1997 117(4) 


reduced to 20) desert bird species for the deserts of Pakistan and India 
(Cowan 1987, 1996), 26 for the rift valley desert of the Gulf of Aden 
and southern Red Sea region (Cowan 1990) and 14 for the 
~ Caspio-Central Asian desert (Cowan 1996), involving 39 different 
species. Each of the species referred to by name in the previous 
paragraph would be a desert bird. 


References: 

Ayyad, M. A. & Ghabbour, S. I. 1986. Hot deserts of Egypt and the Sudan. Pp. 149-202 
in M. Evenari, I. Noy-Meir & D. W. Goodall (eds), Hot Deserts and Arid 
Shrublands, B. Elsevier, Amsterdam. 

Bailey, R. G. 1983. Delineation of ecosystem regions. Environ. Mgmt. 7: 365-373. 
Bailey, R. G. 1989. Explanatory supplement to ecoregions map of the continents. 
Environ. Conserv. 16: 307-309, 310 (map). mo af 
Beard, J. S. 1980. The physiognomic approach. Pp. 33-64 in R. H. Whittaker (ed.), 

Classification of Plant Communities. Junk, The Hague. 

Bradshaw, S. D. 1988. Desert reptiles: a case of adaptation or pre-adaptation? 7. Arid 
Envts. 14: 155-174. 

Casselton, P. J. 1984. Breeding birds. Pp. 229-240 in J. L. Cloudsley-Thompson (ed.), 
Sahara Desert. Pergamon, Oxford. 

Cowan, P. J. 1985. Birds in the central Fezzan, Libya, 1982-1983. Gerfaut 75: 211-218. 

Cowan, P. J. 1987. Birds in the deserts of Pakistan and India: a preliminary review. Bull. 
Ornithol. Soc. Middle East 19: 4-16. 

Cowan, P. J. 1990. Desert birds and the avifauna of the rift valley desert of the Gulf of 
Aden and southern Red Sea region. 7. Arid Envts. 19: 125-131. 

Cowan, P. J. 1996. Desert birds of the Caspio-Central Asian desert. Glob. Ecol. Biogeogr. 
Letts. 5: 18-22. 

Evenari, M. 1985. The desert environment. Pp. 1-22 in M. Evenari, I. Noy-Meir & 
D. W. Goodall (eds), Hot Deserts and Arid Shrublands, A. Elsevier, Amsterdam. 

Jennings, M. C. 1980. Breeding birds in central Arabia. Sandgrouse 1: 71-81. 

Le Houérou, H. N. 1977. The nature and causes of desertization. Pp. 17-38 in M. H. 
Glantz (ed.), Desertification. Westview, Boulder, CO. 

Le Houérou, H. N. 1979. North Africa. Pp. 83-107 in D. W. Goodall & R. A. Perry 
(eds), Arid-land Ecosystems: structure, functioning and management. Vol. 1. 
Cambridge Univ. Press. 

Lewington, I., Alstrém, P. & Colston, P. 1991. A Field Guide to the Rare Birds of Britain 
and Europe. Harper Collins, London. 

Logan, R. F. 1968. Causes, climates, and distribution of deserts. Pp. 21-50 in G. W. 
Brown (ed.), Desert Biology. Vol. 1. Academic Press. 

Lowe, C. H. 1968. Fauna of desert environments. Pp. 567—645 in W. G. McGinnies, 
B. J. Goldman & P. Paylore (eds), Deserts of the World. Univ. of Arizona Press. 
Maclean, G. L. 1976. Arid-zone ornithology in Africa and South America. Proc. Int. 

Orn. Congr. 16: 468-480. 

Maclean, G. L. 1996. Ecophysiology of Desert Birds. Springer, Berlin. 

McGinnies, W. G., Goldman, B. J. & Paylore, P. (eds) 1968: Deserts of the World. Univ. 
of Arizona Press, Tucson. 

Miller, A. H. 1963. Desert adaptations in birds. Proc. Int. Orn. Congr. 13: 666-674 

Monod, T. 1986. The Sahel zone north of the equator. Pp. 203-243 in M. Evenari, I. 
Noy-Meir & D. W. Goodall (eds), Hot Deserts and Arid Shrublands, B. Elsevier, 
Amsterdam. 

Oberlander, T. M. 1994. Global deserts: a geomorphic comparison. Pp. 13-35 m A. D. 
Abrahams & A. J. Parsons (eds), Geomorphology of Desert Environments. Chapman & 
Hall, London. 

Richardson, C. 1990. The Birds of the United Arab Emirates. Hobby, Warrington, U.K. 

Schmidt, R. H. 1979. A climatic delineation of the ‘real’ Chihuahuan Desert. ¥. Arid 
Envts. 2: 243-250. 

Schmidt, R. H. 1989. The arid zones of Mexico: climatic extremes and conceptualization 
of the Sonoran Desert. 7. Arid Envts. 16: 241-256. 

Thomas, D. H. 1984a. Adaptations of desert birds: sandgrouse (Pteroclididae) as highly 
successful inhabitants of Afro-Asian arid lands. 7. Arid Envts. 7: 157-181. 


P. #. Cowan 303 Bull. B.O.C. 1997 117(4) 


Thomas, D. H. 1984b. Sandgrouse as models of avian adaptations to deserts. S. Afr. ¥. 
Zool. 19: 113-120. 

Thomas, D. H. & Robin, A. P. 1977. Comparative studies of thermoregulatory and 
osmoregulatory behaviour and physiology of five species of sandgrouse (Aves: 
Pteroclididae) in Morocco. ¥. Zool. 183: 229-249. 

UNESCO. 1979. Map of the world distribution of arid regions. MAB Tech. Notes 7. 
UNESCO, Paris. 

Verstraete, M. M. & Schwartz, S. A. 1991. Desertification and global change. Vegetatio 
91: 3-13. 

Webster, M. D. 1991. Behavioral and physiological adaptations of birds to hot desert 
climates. Proc. Int. Orn. Congr. 20(3): 1765-1776. 

Whittaker, R. H. 1975. Communities and Ecosystems. 2nd ed. Macmillan, New York. 

Wiens, J. A. 1991. The ecology of desert birds. Pp. 278-310 in G. A. Polis (ed.), The 
Ecology of Desert Communities. Univ. of Arizona Press. 

World Conservation Monitoring Centre. 1992. Global Biodiversity. Chapman & Hall, 
London. 


Address: Dr P. J. Cowan, Department of Zoology, University of Kuwait, P.O. Box 5969, 
Safat 13060, Kuwait. 


© British Ornithologists’ Club 1997 


P. Gregory 304 Bull. B.O.C. 1997 117(4) 


Range extensions and unusual sightings from 
Western Province, Papua New Guinea 


by Phil Gregory 
Received 5 December 1996 


The birds of the Tabubil area of the Western Province of Papua New 
Guinea have now been documented by the papers written by Bell 
(1969), Coates & Lindgren (1978), Murray (1988a) and Gregory 
(1995d). The latter added over 75 species to the previous total 
recorded. ‘The status of some species is altering as habitats change due 
to settlement, and a number of species such as Pied Chat Saxicola 
caprata and Black-headed Whistler Pachycephala monacha have 
colonised new areas of secondary growth. Losses have been few, with 
only the Brown Quail Coturmix australis and Tawny Grassbird 
Megalurus timoriensis seemingly gone, the former due to the loss of its 
grassland habitat at what is now the mine site. 

‘The additions since Gregory (1995d) are summarised below and now 
bring the Ok Tedi area total up to some 329 species, a very high total 
given the primarily forest habitats and absence of extensive wetlands or 
coastline. In addition, distributional information from the remote 
Kiunga and Obo (Middle Fly river region) areas of Western Province is 
given in separate sections. 

For the present purpose, the Ok 'Tedi area may be defined as follows: 
the valley of the Ok Tedi extending south to km 90 on the Kiunga road 
and northwest as far as Mt. Binnie inside the mine site itself. The 
Hindenburg Walls (actually some three such structures, spectacular 
knife-edge limestone escarpments forming part of the foothills of the 
Hindenburg Range) form a natural boundary to the north, and the 
survey area extends west as far as the Lukwi valley beyond the village 
of Ok Ma. The total area is approximately 500 km’. 


Additions and corrections for the Ok Tedi area 
PEACEFUL DOVE Geopelia striata 


Singles in T'abubil town in February and March 1996 represent a 
large northward range extension from the southern Trans-Fly. Suitable 
habitat appears available for colonization as the area is opened up. 


THREE-TOED SWIFTLET Collocalia papuensis 


A single specimen captured at Luplupwintem Cave just outside the 
Ok Tedi survey area in April 1993 was the first such for PNG 
(Rowland 1994). Conceivably much overlooked. There were some 
Collocalia swiftlets flying about in front of thunderheads at km 9 on the 
Ok Ma road in October 1995 with several birds looking like something 
different. They were quite large, with an extremely fast flight more like 
an Apus than the usual Collocalia ‘dithering’, spiky winged and with a 
slightly forked tail, brownish plumage with a prominent silvery chin 


P. Gregory 305 Bull. B.O.C. 1997 117(4) 


and throat which contrasted sharply with the dull underparts. I think 
these are likely to be Three-toed Swiftlet Collocalia papuensis, but a 
specimen would be necessary for certainty. 


GREATER MELAMPITTA Melampitta lugens 

Discovered on Mt. Robinson in forest with no obvious karst 
formations at 1000 m on 31 October 1992, with at least 2 birds calling 
in one area (R. Rowland and P. Gregory). Also heard at two other sites 
on Mt. Robinson at about 1100 m, from Dablin Creek at 750m and 
from km 7 and 9 on the Ok Ma road at 750m where it occurs in the 
more usual limestone karst country. 


GRAY’S GRASSHOPPER WARBLER Locustella gray 

A single L. grayi was found in undergrowth at km 9 on the Ok Ma 
road in October 1995. A probable in March 1992 in similar secondary 
scrub habitat in the town is suggestive of a passage migrant. 


LEMON-BELLIED FLYCATCHER Microeca flavigaster 

Uncommon. A single bird at 700 m at Dablin Creek in early 1993 
was unexpected but the species is now regularly found there, with a 
maximum of 5 birds (probably a family party). Originally thought to be 
Olive Flycatchers M. flavovirescens and listed as such in Gregory 
(1995d) but subsequent views have permitted a re-identification. The 
form here is bright yellow beneath with dark legs and a short tail. 


GREY SHRIKE-THRUSH Colluricincla harmonica 


A single in Tabubil in March 1996 was an unexpected addition, 
maybe a wanderer from the ‘Trans-Fly. 


GREY-STREAKED FLYCATCHER Muscicapa grisetsticta 

Vagrant, but may be more regular than supposed. The third and 
fourth records for Papua New Guinea came from forest clearing edge at 
Ok Menga on 3 November 1991 and 22 November 1992. A spring bird 
was in T’abubil on 15 March 1996, suggesting that the species may be 
regular in the area as it is quite frequent in Irian Jaya during the 
northern winter. 


PAPUAN FLOWERPECKER Dicaeum pectorale 


A common canopy-dwelling species up to about 1500 m locally, often 
seen in pairs. Omitted in error from Gregory (1995d) but previously 
recorded by Murray (1988) and Coates & Lindgren (1978). 


A note on two Ok Tedi mysteries 


Melanocharis sp. 

A female Melanocharis recorded as Obscure Berrypecker M. 
arfakiana by Coates & Lindgren (1978) was seen in hill forest at 640 m 
on 8 February 1978. The yellow pectoral tufts and some orange on the 
gape or bill base suggested an immature bird. Murray (1988a) reported 
a similar yellow tufted bird at the plateau edge on 17 April 1987. 


P. Gregory 306 Bull. B.O.C. 1997 117(4) 


At Dablin Creek on 1 October 1992, C. Eastwood and I found two 
small canopy-haunting Melanocharis-type berrypeckers with yellow 
pectoral tufts and orange bills, but also with faint dark streaks 
underneath (Gregory 1993a). In July 1993 orange-billed yellow-tufted 
birds without streaks were common in the same area, haunting the 
canopy and having a high pitched buzzy song identical to unidentified 
berrypeckers discovered at Boana on the southern Huon by a Field 
Guides Inc. tour group in 1991. These Boana birds have yellow 
pectoral tufts, but also an eye ring, which is lacking in the Tabubil form 
(B. Whitney and J. Pierson pers. comm.). 

Clearly much work needs to be done to reveal the identity of these 
forms, which may be referable to M. arfakiana, or represent an 
undescribed taxon. Two individuals’ were collected under permit in 
July 1994 by Whitney and are being studied in the U.S.A. The form is 
quite frequent at heights of c. 750-1000 m in the area, now being 
known from the Ok Ma road and Mt. Robinson as well. 

It is also worth recalling that Coates & Lindgren (1978) reported a 
drab olive Melanocharis of the striativentris/longicauda type at 2220 m 
on Mt. Binnie in moss forest, but with strong black streaking down the 
sides of the abdomen and white pectoral tufts. There have been no 
further sightings as yet and its identity remains unknown. 


RED-BREASTED PIGMY-PARROT Micropsitta bruijnit 

A form resembling Red-breasted Pigmy-parrot, but with a 
distinctive yellowish cap, is common in the area. This form was first 
recorded by Murray (Murray 1988b) on Mt. Binnie on 20 September 
1987 and was rediscovered by G. Johnston and myself at Dablin 
Creek on 29 November 1991 (Gregory 1994). There have been 
frequent sightings. since with 40 flying N at Ok Menga at 750m on 
4 October 1992; also 220 flying N there on 25 October 1992; 45 on 
8 November 1992, and an unprecedented total of 650 heading NE 
on 22 November 1992 between 16.00 and 16.45 hrs, in flocks of 50-60 
birds with one flock of over 100. Flocks of 20-30 are common at Dablin 
Creek. A pair were excavating a chamber in a stump at Townsville at 
1650 m in September 1992 (photograph of the male in Gregory 1995d). 

Reference to museum specimens in Port Moresby, Sydney and 
Hawaii by G. Johnston suggests some variation in the colouration of 
males of this species, and the local birds are likely to be within this 
continuum rather than being a new subspecies. 

The Yellow-capped Pigmy-parrot WM. kezensis is unrecorded from the 
area (or confused with the other form) but is possible lower down the 
valley. 


Further notes on the plumage phases of the Grey-headed Goshawk 

I recorded a previously undescribed melanistic morph of the 
Grey-headed Goshawk Accipiter poliocephalus in the Ok Ma area in 
1992 (Gregory 1995c). A normal phase adult, and an adult of a hitherto 
undescribed greyish morph, were found at Ok Menga, Western 
Province, on 21 September 1996. The latter was twice seen well, 
perched in trees, and watched through a’ X 30 telescope on my second 


P. Gregory 307 Bull. B.O.C. 1997 117(4) 


view. It showed whitish underwing coverts in flight, like the normal 
phase adult seen earlier. Its plumage was otherwise entirely mid-grey, 
including the head and underparts, with no streaks or bars, quite unlike 
the usual pale headed appearance. The legs were the normal bright 
pink, and the cere was also bright pink, extending back to the dark eye 
which had a reddish ring around it joined to the pink loral patch. In its 
head pattern the bird was curiously reminiscent of a Gabar Goshawk 
Micronisus gabar. These New Guinea accipiters remain poorly known, 
particularly their immature plumages, and the Grey-headed Goshawk 
is clearly polymorphic in this area. 


Distribution data from the Kiunga area 


The Kiunga area, along the upper Fly River some 140 km south of 
Tabubil, is well-known for a number of rare New Guinea species, and 
some recent records are given below. New road and _ forestry 
developments look set to cause significant changes in the near future. 


NEW GUINEA FLIGHTLESS RAIL Megacrex inepta 


Local informants report this species as being not rare in dense sago 
swamps. A captive individual with a single leg was in Kiunga in August 
1994. 


SOUTHERN CROWNED PIGEON Goura scheepmakeri 

Still to be found in small parties in the monsoon forest up-river, 
though extirpated from the immediate vicinity of the town. Birds are 
quite frequently traded in local markets. 


VULTURINE PARROT Psittrichas fulgidus 

This highly-prized species, classified as Vulnerable by BirdLife 
International, is occasionally seen flying over the river, or along 
Magazine Road north of the town. 


LARGE FIG-PARROT Psittaculirostris desmarestit 


This spectacular and uncommon species is sometimes seen from the 
river, usually in pairs. 


LESSER PARADISE-KINGFISHER Tanysiptera hydrocharis 

Reported in October 1992 from beyond Magazine Road (A. Richards 
and R. Rowland in /itt.) and April 1994 about 20 minutes up-river (I. 
Richardson in /itt.); sympatric with the Common Paradise-Kingfisher 
T. galatea which is frequent here. A single bird was also found in 
August 1996, in slightly drier forest than the Common Paradise- 
Kingfisher (B. Whitney pers. comm.). 


LONG-BILLED CUCKOO Rhamphomantis megarhynchus 

Recorded on the Field Guide Inc tours in July and August during 
the early years of the decade, and again in 1996, about 15 km north of 
the town on the Drimgas road. This is a significant extension of the 
known range of this rare and cryptic species from its previous sites 


P. Gregory 308 Bull. B.O.C. 1997 117(4) 


in the Middle Sepik district and around the Port Moresby/Central 
Province area. 


WHITE-BELLIED PITOHUI Pitohui incerta 


This rare and enigmatic species occurs up-river from the town in the 
monsoon forest, with brown and black bird parties. It may easily be 
mistaken for a Little Shrike-thrush Colluricincla megarhyncha but is 
stouter, shorter tailed, and pale creamy beneath with fairly obvious 
diffuse darker mottling on the chest. Prominent dark eye with pale 
lores, being almost encircled by pale like an eye-ring. Bill pinkish-horn 
with a darker lower mandible. Legs brownish-pink. One distinctive call 
was a repeated tuc, almost like clicking the tongue, as well as some 
typical ‘pitohui type’ musical calls. The birds kept to the mid-stratum, 
feeding in vine tangles and perching on tree trunks in a manner 
reminiscent of Phyllastrephus bulbuls. 


BROAD-BILLED FAIRY-WREN Malurus grayi campbelli 


Discovered by Palliser (1988) in swamp forest between the Kiunga 
airstrip and the river. ‘There are no subsequent Kiunga records, but it 
was found in a peculiarly difficult and inaccessible habitat. This may be 
the only field observation of this form, the Mount Bosavi (extreme 
southwest of Southern Highlands Province) birds all being trapped and 
then described as a new species M. campbelli (Schodde & Weatherly 
1982). Following further research and specimens, it was subsequently 
shown by LeCroy & Diamond (1995) to be a subspecies of the 
Broad-billed Fairy-wren M. grayt. 


BANDED YELLOW ROBIN Poecilodryas placens 


Reported from monsoon forest a short way up-river in April 1994 (I. 
Richardson in Ilitt.). Confirmation is desirable; this would be a 
significant but not unlikely range extension, as the species is previously 
known from Mount Bosavi. 


WHITE-SPOTTED MUNIA Lonchura leucosticta 

Reported by Palliser (1988) from the airstrip vicinity, and found in 
April 1994 at Rumgenai some 27 km north of the town. A Trans-Fly 
species that is on the edge of its range here, Trans-Fly being the lower 
and middle sections of the Fly river catchment. 


CRIMSON FINCH Neochmia phaeton 


Two birds with a White-spotted Munia in kunai grass at Rumgenai 
in April 1994 were a long way from the known range in the middle 
Fly River. 


YELLOW-EYED STARLING A plonis mystacea 


Occurs in small numbers with flocks of Metallic Starlings A. 
metallica along the river. The nasal tuft is a useful field character in 
flight. 


P. Gregory 309 Bull. B.O.C. 1997 117(4) 


FLAME BOWERBIRD Sericulus (aureus) ardens 


The southern form S. (aureus) ardens is still to be found along 
Magazine road north of the town, where the birds fly over at canopy 
height of an early morning; also quite often seen flying over the 
Boystown Road. Skins of males are occasionally seen dangling from the 
driving mirrors of vehicles, where they are a local version of fuzzy dice 
and clearly an object of status! 


GREATER and RAGGIANA BIRDS-OF-PARADISE Paradisaea apoda and 
P. raggiana 

The Raggiana seems to be the species usually seen along the Fly 
River itself in this district, whilst the Greater occupies the areas further 
away. Hybrids would be extremely difficult to detect, and whether the 
two species do hybridize here is still unknown. One lek of Greater 
Birds has been in use for over two decades. 


Observations from Obo 


Obo is a small airstrip and distribution station sited on the Fly River 
just north of the Strickland confluence, lying in the middle-Fly 
wetlands about 200 river miles inland. Ok Tedi Mining Ltd sponsored 
some surveys in 1994-95 for the Asian Wetland Bureau, and the 
records here were made whilst a member of that team, unless otherwise 
stated. Very little has been published about the area. 


LITTLE BITTERN /xobrychus minutus 


Jaensch (1995) recorded four calling birds in June 1994, flushing a 
probable pair—which strongly suggested that breeding occurs in these 
vast wetlands. Various other individuals were seen in December 1994 
and April 1995. It can be assumed that a resident breeding population 
is present, which may perhaps be seasonally augmented by Australian 
migrants. 


GLOSSY IBIS Plegadis falcinellus 

Counts of 973 at dusk at Lake Daviumbu on 2 December, and 400+ 
at Lake Ambuve on 4 December show the significance of the area for 
this species during the austral summer. None were seen in April 1995. 


MAGPIE GOOSE Anseranas semipalmatus 

The area holds good numbers during the austral summer, with 250 at 
Lake Pangua on 2 December and 150 at Lake Owa on 3 December 
1994. Much smaller numbers were present in April 1995, and nests 
were found. 


GARGANEY Anas querquedula 


4 at Lake Pangua 2 December, 12 at Lake Owa 3 December, 38 Lake 
Ambuve 4 December 1994. The species is sparse in Papua New Guinea 
and this may be a significant wintering area. 


P. Gregory 310 Bull. B.O.C. 1997 117(4) 


YELLOW WAGTAIL Motacilla flava 

The marshy edges of ox-bows supported good numbers of birds, 
with.10 at Lake Pangua on 2 December, 45 at Lake Owa on 
3 December and 10 at Lake Ambuve on 4 December. The area is likely 
to be a significant wintering ground, for a species thought.to be very 
scarce in Papua New Guinea. 


RESTLESS FLYCATCHER Myzagra inquieta 

The species was previously only known from the southern Bensbach 
River area in Papua New Guinea, so singles near Komavai village, Lake 
Pangua on 26 April and at Obo station on 27 April 1995, and a pair 
found nesting on the roof beam of an abandoned hut at Lake Ambuve 
in April 1995, are noteworthy range extensions, several hundred 
kilometres further inland. The nest was a neat cup of dried grass and 
fine woody material containing a single whitish egg with dark spots. 
Apparently the first instance of nesting proven for New Guinea. 


GREAT REED WARBLER Acrocephalus arundinaceus 

Usually considered to be a vagrant to the Australasian region. One 
was singing in reeds along the channel to the main river at Lake 
Ambuve on 4 December, readily recognised by the distinctive harsh 
song, large size and streaked breast. May be a far more regular visitor 
than previously realised. 


FLY RIVER GRASSBIRD Megalurus albolimbatus 

Two at Lake Daviumbu on 2 December, the type locality for the 
species; one at Lake Owa on 3 December. Recorded in April from Lake 
Pangua, Lake Owa and Lake Daviumbu, in floating Echinochloa and 
Leersia sp. grass mats, not Cyperus sedge beds as is the habitat at 
Bensbach. ‘The species has a distinctive whistled call, a quiet, rising 
zeeee, and may be distinguished from Tawny Grassbird M. timoriensis 
by the prominent white edges to the black tertials, a gently rounded 
rather than ragged tail, orangish rump, whitish supercilium and 
underparts and an unstreaked orangish crown. 

Lake Owa and Lake Pangua are new localities for the species which is 
a Papua New Guinea endemic previously known from Lake Daviumbu 
and the middle Bensbach river. ‘The abundance of suitable habitat in 
the middle Fly suggests that this easily overlooked species may be more 
widely distributed than currently recognised and may occur across the 
border in West Irian. 


BLACK MANNIKIN Lonchura stygia 

Not recorded during December 1994, and only found at Lake Owa in 
April 1995, despite extensive searches at other apparently similar sites 
during both expeditions. We saw a total of over 30 individuals in small 
flocks, most with flocks of Grey-crowned Mannikin Lonchura 
nevermanni, which was widespread in the area in both December and 
April. Lonchura stygia was invariably seen at rest in floating or 
emergent grass, especially Oryza and Echinochloa spp. in full seed head, 
feeding on Echinochloa seeds like its congener. 


P. Gregory 311 Bull. B.O.C. 1997 117(4) 


Birds were in a variety of plumages (Gregory 1995b), and are clearly 
very scarce in the Obo area where the species is presumably nomadic, 
dependent on supplies of seeds. The species is only otherwise known 
in Papua New Guinea from Lake Daviumbu, being endemic to the 
western Trans-Fly. We did not find it at this site during our visits and 
our observations suggest it is rather rare. 


Acknowledgements 


I would like to thank Chris Eastwood and Dr Ian Burrows for their records, and K. 
David Bishop, Roger Jaensch and Bret Whitney for access to trip reports and verbal 
observations. Dr J. M. Diamond provided valuable comments and data about the Greater 
Melampitta. Ok Tedi Mining Ltd. must be thanked for access to mine and drill sites and 
logistic support, specifically the Environment Department and Ian Wood, Dr Murray 
Eagle and Dr Andrew Storey. I also thank the various local landowners, particularly 
Borok Pitalok of Wangbin, for allowing me access to their customary land. 


References: 

Bell, H. L. 1969. Field notes on the birds of the Ok Tedi River drainage, New Guinea. 
Emu 69: 193-211. 

Coates, B. J. & Lindgren, E. 1978. Ok Tedi Birds. Ok Tedi Environmental Task Force, 
Ok Tedi Development Co. 

Gregory, P. 1993b. An unidentified berrypecker Melanocharis sp. from the Ok Tedi area. 
Muruk 6: 33. 

Gregory, P. 1994. Further records of an unidentified pigmy-parrot Micropsitta sp. from 
the Ok Tedi area. Muruk 6: 11-12. 

Gregory, P. 1995a. Further studies of the birds of the Ok Tedi area, Western Province, 
Papua New Guinea. Muruk 7: 1-38. 

Gregory, P. 1995b. Notes on the Black Mannikin (Lonchura stygia) and other mannikins 
(Lonchura spp.) at Lake Owa, Middle-Fly River. Muruk 7: 119-120. 

Gregory, P. 1995c. An undescribed plumage morph of the Grey-headed Goshawk 
(Accipiter poliocephalus). Muruk 7: 116. 

Gregory, P. 1995d. The Birds of the Ok Tedi Area. Ok Tedi Environment Dept. 

Jaensch, R. 1995. Little Bitterns Zxobrychus minutus) in the Middle-Fly wetlands. Muruk 
7: 117-118. 

LeCroy, M. & Diamond, J. 1995. Plumage variation in the Broad-billed Fairy-wren 
Malurus grayi. Emu 95: 185-193. 

Murray, A. 1988a. A study of the birds of the Tabubil region, Western Province, Papua 
New Guinea. Muruk 3: 89-117. 

Murray, A. 1988b. A possible new pigmy-parrot Micropsitta. Muruk 3: 118. 

Palliser, T. 1988. Papua New Guinea: Report on a birding trip July to December 1987. 
Privately published. 

Rowland, P. 1994. A new altitude record and range extension for the Three-toed Swiftlet 
Collocalia papuensis. Muruk 6: 10. 

Schodde, R. & Weatherly, R. G. 1982. Campbell’s Fairy-wren Malurus campbelli, a new 
species from New Guinea. Emu 82: 308-309. 


Address: P. Gregory, P.O. Box 69, Tabubil, Western Province, Papua New Guinea. 


© British Ornithologists’ Club 1997 


In Brief 312 Bull. B.O.C. 1997 117(4) 
IN BRIEF 


A NEW BREEDING AREA FOR THE SONG THRUSH 
TURDUS PHILOMELOS IN NORTH PORTUGAL 


The Song Thrush is a partially migratory Palaearctic species which 
breeds over a vast area from Western Europe to Central Asia. The 
Iberian Peninsula constitutes an important wintering area for this 
species. The movements of the partially migratory populations may be 
affected by poor weather (Santos 1982, 1985). 

The historical evidence concerning the possible breeding of this 
species in Portugal is mainly restricted to the Portuguese national 
breeding Atlas, which refers to possible nesting in the Coroa Hills 
(northeast Portugal) (Rufino 1989). Earlier, Tait (1924) mentioned 
nesting of this species in the Cabrera Hills (Puebla de Sanabria), on the 
Spanish side of the boundary of the Montesinho Natural Park 
(northeast Portugal). More recently, its breeding was confirmed in 
Portugal for the first time. In 1991, a nest was discovered in 
Peneda-Gerés National Park (northwest Portugal) (Santarém 1991, 
Pimenta & Santarém 1996). 

In field surveys for the breeding Atlas of the Montesinho Natural 
Park (north Bragan¢a) (1992-1995) and the breeding Atlas of Nogueira 
Hills (southwest Braganca) (1995-1996) the Song ‘Thrush was 


North Spain 


North-west Portugal 


Central Spain 


Central Portugal 


@ Possible breeding 
@ Probable breeding 


@ Confirmed breeding 


Figure 1. Distribution map of the Song Thrush Turdus i ag breeding area in 
northeast Portugal, based on 10 x 10 UTM squares. 


In Brief 313 Bull. B.O.C. 1997 117(4) 


TABLE 1 
Location. of confirmed, probable, or possible breeding records of Song Thrushes in 
northeast Portugal 


10x 10 UTM Breeding 
Place Year squares Habitat status 
Montesinho Hills (Montesinho) 1992 PG 84 Oak woodland Possible 
Montesinho Hills (Vilarinho) 1994 PG 74 Riparian vegetation Possible 
Rabal 1992 PG 83 Riparian vegetation Confirmed 
Soeira 1993 PGs Riparian vegetation Probable 
Vinhais 193 PG’63 Riparian vegetation Probable 
Cobelas 1994 PCs Riparian vegetation Possible 
Moimenta 1295 PG 64 Agro-forestry Possible 
Gestosa 1994 PGi63 Agro-forestry Possible 
Nogueira Hills 1995 PG 82 Riparian vegetation Probable 
Nogueira Hills 1996 PG 81 Riparian vegetation Confirmed 


detectedin nine 10 x 10 km UTM squares. These data represent the 
first breeding evidence of this species in the northeast of the country, 
indicating the existence of a second and larger nucleus in northern 
Portugal. 

It occurs mostly in riparian vegetation consisting of Common Alder 
Alnus glutinosa, Narrow-leaved Ash Fraxinus angustifolia and poplars, 
or White Poplar Populus alba and Black Poplar P. nigra associated with 
moors and highland pastures. It also occurs in oak woods and mixed 
deciduous woods formed mainly by Pyrenean Oak Quercus pyrenaica 
and Birch Betula sp. (see Table 1). 

The apparent absence of this species in northern Portugal in the 
past was mainly due to an insufficient coverage of the more remote 
areas and did not _ necessarily reflect its real distribution. 
Nevertheless, it is believed that in recent years this species’ breeding 
distribution has expanded southwards to certain areas in the north of 
Portugal, resulting in the distribution presented here for the 
northeast (Fig. 1). To support this idea some local hunters report 
that this thrush has only bred in Nogueira Hills for the last five 
years. It should be noted that the distribution presented may still be 
somewhat incomplete due to difficulties in detecting Song Thrushes 
breeding at low densities. 


References: 

Pimenta, M. & Santarém, M. L. 1996. Atlas das Aves do Parque Nacional da 
Peneda-Gerés. Edic¢ao do Instituto da Conservacao da Natureza e do Parque Nacional 
da Peneda-Gerés, Porto. 

Rufino, R. (coordinator ) 1989. Atlas das Aves que nidificam em Portugal Continental. 
Servico Nacional de Parques, Reservas e Conservacao da Natureza, Lisboa. 

Santarém, M. L. 1991. Primeiros dados sobre a nidificagéo do Tordo-comun Turdus 
philomelos em Portugal. Airo 2: 52-53. 

Santos, T. 1982. Migracién y invernada de Zorzales y Mailros (genero Turdus) en la 
Peninsula Ibérica. Tesis Doctoral. Universidad Complutense de Madrid, Madrid. 


In Brief 314 Bull. B.O.C. 1997 117(4) 


Santos, T. 1985. Estudio sobre la biologia migratoria de la Tribu Turdini (Aves) en 
Espana. SEO-ICONA. Monografia 39, Madrid. 
Tait, W. 1924. The Birds of Portugal. Witherby, London. 


Av. Dr. Mario Moutinho L-1733, 10E, LUIS MIGUEL REINO 
1400 Lisbon, 

Portugal. 

Quinta da Olaia, DOMINGOS PATACHO 
2490 Ourém, | 
Portugal. 13 November 1996 


FIRST DESCRIPTION OF THE NEST OF THE SLENDER-BILLED FINCH 


The Slender-billed Finch Xenospingus concolor is a poorly known bird 
that inhabits some coastal valleys from central Peru to northern Chile 
(Koepcke 1970, Ridgely & Tudor 1989). It is shy and inconspicuous, 
occurring in natural riparian shrubbery, and its nest has not been 
described (Collar et al. 1992). Here I present observations on two nests 
in southwestern Peru. 

On 18 December 1995, I found an individual building a nest in the 
riparian thickets of La Real (16°07'S, 72°29’W), in the Majes valley at 
an elevation of 550m. Slender-billed Finches occur in the riparian 
thickets (mainly of Tessaria integrifolia and Baccharis sp.) all along 
this valley (Gonzalez & Malaga 1997). The site of the nest was about 
50 m from the river, and on the other side about the same distance 
from extensive rice plantations. A pair of birds was present in the 
vicinity of the site but only one bird, presumably the female, was 
nest-building. During three hours of observations, it twice brought 
twigs to the nest, and twice went to the nest without material, 
apparently to shape the interior. The nest was placed among 
ramificating dead branches of Tessaria with a tangle of many dead 
stems and leaves of Baccharis. It was about 80% complete, 2 m above 
ground and well hidden; globular in shape, about 11 cm in diameter, 
and entirely woven of fine twigs. 

The second nest was found on 2 June 1996, in riparian shrubbery 
between Ocucaje and Callango (14°27'S, 75°38’W), in the Ica valley 
at an elevation of 300 m. It was first located by students taking part 
in a field course on regional wildlife, and shown to me. The 
riparian shrubbery of Ocucaje is drier than that of the Majes valley, 
and interspersed with Baccharis (fruiting at that time), ““Huarango”’ 
trees (Acacia macracantha; with fruit in pods) and introduced 
shrubs of Tamarix. 'The nest was empty and apparently recently 
built, as a pair of Slender-billed Finches were calling persistently 
around it, uttering high-pitched cheeping calls which at first led us 
to suppose that there were nestlings. in the nest. This nest was 
woven with dry leaves of “‘cana brava’ (Gynerium sagittatum, a 
large grass) and was oval-shaped, with the entrance high up on one 
side; height 12-13 cm, depth 8cm, width of nest walls 2.2 cm, 
entrance diameter 4cm. It was 1.5 m above ground, in a tangle of 


In Brief 315 Bull. B.O.C. 1997 117(4) 


small twigs between the branches of a small Acacia, with an 
impenetrable tangle of ‘“‘cana brava’ behind. It was collected and is 
now in the collection of the Departamento de Ornitologia, Museo 
de Historia Natural of the Universidad Nacional Mayor de San 
Marcos, Lima. 


This discovery was one of the objectives in a research project on the ecology of the 
Slender-billed Finch, sponsored by the American Bird Conservancy. I wish to thank 
Biol. Victor Velasquez for inviting me to Ocucaje and the students that helped me to 
discover the nest: Emilio Ayquipa, Jessica Caballero, Erika Carnero, Jorge Espinoza, 
Alipio Baldeon, all from the Facultad de Ciencias Biologicas y Quimicas of the 
Universidad Nacional de Ica. Irma Franke (Museo de Historia Natural, UNMSM, 
Lima) and Manuel Plenge helped with the MS. 


References: 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, N. G., Parker, 
T. A. & Wege D. C. 1992. Threatened Birds of the Americas. The ICBP/IUCN Red 
Data Book. 3rd edn, Part 2. ICBP, Cambridge, U.K. 

Koepcke, M. 1970. The Birds of the Department of Lima. Livingston Publ. Co., 
Wynnewood. 

Gonalez, O. & Malaga, E. 1997. Distribucion de aves en el Valle de Majes, Arequipa, 
Peru. Orn. Neotropical 8: 57-69. 

Ridgely, R. & Tudor, G. 1989. Birds of South America. Vol. 1. The Oscine Passerines. 
Univ. Texas Press. 


Gomez del Carpio 135, OSCAR GONZALEZ M. 
Barrio Médico, 

Lima 34, 

Peru. 20 December 1996 


A SPECIMEN OF HIRUNDO ABYSSINICA FROM MADAGASCAR 


In the course of a revision of the bird collection in the Museum of 
Natural History of Geneva, Switzerland, I came across a specimen of 
Hirundo abyssinica which appears to be the first record for Madagascar. 
The specimen (MHNG 835.83) was purchased in September 1925 
from a certain Mr Chauvin, along with six other Malagasy species. The 
shipment was dispatched in part from the Académie Malgache. The 
specimen was collected in January 1925 near Lake Alaotra, northeast of 
Antananarivo. In eastern Africa, the species breeds from Ethiopia to 
the Cape and is partly migratory, so its occurrence in Madagascar as an 
occasional, if not regular, visitor is not surprising. Dr Francois Baud, 
Curator, kindly provided further details on the origin of the specimen. 


rue de Prévent, MICHEL DESFAYES 
1926 Fully, 
Switzerland. 30 December 1996 


FIRST RECORD OF SWAINSON‘'S HAWK BUTEO SWAINSONI FOR THE 
WEST INDIES 


On the afternoon of 22 April 1996, c. 3km south of Monte Christi, in 
the northwest Dominican Republic, the authors were driving through 


In Brief 316 Bull. B.O.C. 1997 117(4) 


an area of dry scrub when a medium-sized Buteo was noticed soaring 
near the road. We stopped and watched the bird for approximately five 
minutes at ranges down to c. 100 metres before it flew west and out of 
sight. We identified it as a Swainson’s Hawk Buteo swainsoni, a species 
both GMK and CGB had seen previously in the U.S.A. and Canada. 

The bird was identified as a light morph adult and the following 
details were recorded. Approximately the same size as Red-tailed Hawk 
Buteo jamaicensis, but with marginally longer, more pointed wings and 
narrower hand than that species. The head was dark brown with 
whitish throat that contrasted with the dark brown upper breast. The 
lower breast, belly and ventral area were white. The tail was dark. The 
underwings had whitish underwing coverts contrasting with darker 
remiges recalling Booted Eagle Hieraaetus pennatus, although the 
remiges were not as uniformly dark as that species. The upperparts 
appeared uniformly dark brown. 

This is the first documented sighting of Swainson’s Hawk in the 
Dominican Republic. Stockton de Dod (1981) and Wetmore & Swales 
(1931) list no records for the island of Hispaniola and there are no 
records from elsewhere in the West Indies. The species breeds in 
prairie and open areas of western North America from southern 
Canada south to Texas and central California. It migrates south to 
winter in South America and thus its appearance in the West Indies is 
not unexpected. 


References: 

Stockton de Dod, A. 1981. Guia de campo para las aves de la Republica Dominicana. 
Editora Horizontes de America, Santo Domingo. 

Wetmore, A. & Swales, B. H. 1931. The birds of Haiti and the Dominican Republic. 
Bull. U.S. Natn. Mus. 155. 


6 Collet Walk, CHRIS G. BRADSHAW 
Parkwood, 
Gillingham, Kent ME8 9QL. 


6 Connaught Road, GUY M. KIRWAN 


Norwich, 
Norfolk NR2 3BP. 


5a West Parade, ROBERT S. R. WILLIAMS 


Norwich, 
Norfolk NR2 3DN. 15 March 1997 


NOTES ON THE BLACK-BACKED TANAGER TANGARA PERUVIANA 
(DESMAREST, 1806) 


Tangara peruviana has a limited range in southeast Brazil, from 
Espirito Santo to Rio Grande do Sul (Meyer de Schauensee 1970, 
Argel de Oliveira et al. 1993). It has been considered conspecific with, 
and a colour morph of, the more widespread T. preciosa (Sick 1988), 
but is now recognised, though somewhat tentatively, as a separate 


In Brief 317 Bull. B.O.C. 1997 117(4) 


species (Sibley & Monroe 1990). Collar et al. (1992) consider it 
threatened, as most recent records are from habitats in SA0 Paulo and 
Rio de Janeiro states that are rapidly being destroyed. Gonzaga (1987) 
comments on the lack of information about this bird. 

We present here data collected at Pontal do Sul (25°32’S, 49°27’W) 
and Ilha do Mel (25°30’S, 48°21’W), Paranagua, Parana. During field 
work from April 1989 to October 1991 we obtained some records 
of Black-backed Tanagers, and in the course of a_ bird-banding 
programme from November 1991 to July 1994 we captured a number 
of individuals. These we marked with CEMAVE/IBAMA numbered 
rings, noting their weight and wing, tail and bill measurements, and 
details of moult. Faeces deposited in the bags in which they were 
brought back to the camp were preserved in 70% alcohol and later 
examined in the laboratory. 

T. peruviana is one of the commonest tanagers at the two above 
localities; single individuals or groups may often be seen foraging in the 
middle and upper strata of the vegetation. Ilha do Mel and Pontal do 
Sul are covered by the so-called restinga formation, defined by Silva 
et al. (1994) as a mosaic of sandy beaches, dunes, forests, mangroves, 
herbaceous and arboraceous marshes. There are also areas with human 
disturbance. From July 1989 to September 1991 we had 13 records at 
Ilha do Mel, of which 8 were from disturbed areas. From 1992 to 1994 
we caught 24 individuals, 16 at Ilha do Mel (6 in disturbed, 10 in 
undisturbed areas) and 8 at Pontal do Sul in areas with human 
disturbance in contact with mangroves. 

The occurrence of Black-backed Tanagers in areas modified by man 
can be related to the presence of suitable fruit-bearing plants. Species 
of Melastomataceae, favourite food plants for tanagers, are very 
common, and there are also Cecropia sp. (Moraceae), Salacia 
sp. (Celastraceae), Gaylussacia brasiliensis (Ericaceae), Psidium sp. 
(Myrtaceae) and other cultivated plants. The faecal analysis showed a 
predominance of fruits (66.6% by frequency) in their diet, with some 
insects (Hymenoptera 16.6%, Coleoptera 16.6%) and spiders (16.6%). 

Weights and measurements of 15 mist-netted birds were as follows: 
weight, mean 22.16, range 18.5—25.5 g, n=15; wing, mean 75.45, range 
70-80 mm, n=24; tail, mean 48.66, range 44-59 mm, n=21; tarsus, 
mean 19.55, range 16.6—22 mm, n=24; bill length, mean 11.05, range 
8.5-13.6 mm, n=24. Twelve of the birds caught showed some moult of 
body feathers; 3 (2 adults in March, 1 juvenile in February) were 
moulting flight-feathers. 

Black-backed Tanagers were present at Ilha do Mel and Pontal do 
Sul in all seasons. They may be totally sedentary, or perhaps partially 
migratory. Little is known about their possible movements. Ihering 
(1885, in Sick 1981) assumed that they travelled short distances in 
connection with the ripening of fruits; other authors, such as Sick 
(1981), have considered the possibility of large-scale migrations. Data 
from the literature show their presence at different localities in Rio 
Grande do Sul at all seasons (Pinto 1944, Camargo 1962, Sick 1988), 
and in Sao Paulo state in winter and summer (Pinto 1944). Krul & 
Moraes (1993) recorded the species at Curitiba, Parana, in all seasons. 


In Brief 318 Bull. B.O.C. 1997 117(4) 


We provisionally conclude that some populations may opportunistically 
travel for short distances and that others are essentially sedentary. 
Their tolerance of human disturbance.and exploitation of fruit sources 
provided by man justify some optimism for their future survival. 


References: 

Argel de Oliveira, M. M., Scardua, C. E. & Paccagnella, S. G. 1993. Ocorréncia da 
saira-sapucaia (Tangara peruviana) (Passeriformes: Thraupinae) no Estado do 
Espirito Santo. Jn: Congresso Brasileiro de Ornitologia, 3., Pelotas, RS. Resumos. 
Pelotas: Universidade Catolica de Pelotas. 

Camargo, O. R. 1962. Aves sul-riograndenses do Museu de caga e pesca. Pesquisas 14: 
1-67. 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono-Nieto, A., Naranjo, L. G., Parker, 
T. A. III & Wege, D. C. 1992. Threatened Birds of the Americas. 1.C.B.P./1.U.C.N. 
Red Data Book. Vol. 2. International Council for Bird Preservation, Cambridge. 

Gonzaga, L. P. 1987. Aves brasileiras ameacadas de extingao. In: Encontro Nacional de 
Anilhadores de Aves, 2., Rio de Janeiro, RJ, 1986. 

Krul, R. & Moraes, V. S. 1993. Aves do Parque Barigui, Curitiba, Parana. Biotemas 6(2): 
30-41. 

Meyer de Schauensee, R. 1970. A Guide to the Birds of South America. Livingston 
Publishing Co. 

Pinto, O. M. O. 1944. Catdlogo das aves do Brasil. 2. Ordem Passeriformes: Superfamilia 
Tyrannoidea e Subordem Passeres. Secretaria da Agricultura, Industria e Comércio, 
Sao Paulo. 

Sibley, C. G. & Monroe, B. L. Jr. 1990. Distribution and Taxonomy of Birds of the World. 
Yale Univ. Press. 

Sick, H. 1981. Migracdes de aves na América do Sul continental. Publ. Técn. CEMAVE 
2: 1-86. 

Sick, H. 1988. Ornitologia brasileira, uma introdugdo. Vol. 2. Universidade de Brasilia, 
Brasilia. 

Silva, S. M., Britez, R. M., Souza, W. S., Lima, R. X. & Athayde, S. F. 1994. Vegetacao 
e flora da Ilha do Mel. In: S. M. Silva & S. F. Athayde (eds), I/ha do Mel: Historia 
Natural e Conservacao. Univ. Federal do Parana, Curitiba. 


Universidade Estadual de Campinas, VALERIA S. MORAES 
Departamento de Zoologia 

13.081-970 Campinas, 

Sao Paulo, Brazil. 


Universidade Federal do Parana, RICARDO KRUL 
Centro de Estudos do Mar, 

Av. Beira-mar s/n, 

83.255-000 Pontal do Sul, 

Parana, Brazil. 23 November 1995 


Books Received 319 Bull. B.O.C. 1997 117(4) 
BOOKS RECEIVED 


Olsen, K. M. & Larsson, H. 1997. Skuas and Jaegers: a guide to the skuas and jaegers of 
the world. Pp. 190, 13 plates (12 in colour). 156 text-figures (21 in colour). Pica Press. 
ISBN 1-873403-16-1. £24. 25 x 17 cm. 


The main purpose of this richly illustrated book, dealing with only seven species, is to 
provide as complete as possible a guide to their identification—often, under field 
conditions, a notoriously difficult problem. The first 16 pages of text give a general 
overview of the subfamily Stercorariinae; this is followed by the species accounts, which 
go in meticulous detail into plumage and other characters, each account ending with a 
full-page distribution map. The 13 plates by Larsson, each illustrating up to 9 birds 
(mostly in flight, some standing), are excellent, and are supplemented by a large number 
of photographs, many in colour. Its size, though rather large for a field-guide, should be 
no hindrance to the ship-borne seabird-watchers who are most likely to find it useful, or 
even essential. 


Lefranc, N. & Worfolk, T. 1997. Shrikes: a guide to the shrikes of the world. Pp. 192, 16 
colour plates, maps, text-figures. Pica Press. ISBN 1-873403-47-X. £25. 25 x 17 cm. 


With an authoritative text (by Norbert Lefranc) and a superb set of colour plates (by 
Tim Worfolk), this is an attractive addition to the Pica Press family guides. Although it is 
stated that the book is “primarily an identification guide’, the 30-page introductory 
section is of a very high standard, dealing not only with taxonomy and relationships but 
also very thoroughly with feeding habits and many other aspects of behaviour and 
ecology. 


Inskipp, T., Lindsey, N. & Duckworth, W. 1996. An Annotated Checklist for the Birds of 
the Oriental Region. Pp. 224. Oriental Bird Club (c/o The Lodge, Sandy, Bedfordshire 
SG19 2DL, U.K.). ISBN 0-9529545-0-8. £9.95. 24 x 17 cm. 


A very useful list of the 2586 species of birds recorded in the area covered by the 
Oriental Bird Club. It is stressed in the introduction that its aim is to summarise 
published data concerning their systematics and taxonomy (distribution is not dealt 
with), and that it is “‘only as complete as are the available data’. It has clearly been 
thoroughly researched and carefully prepared, with annotations mentioning all 
unresolved points of taxonomy backed up by an impressive list of 814 references. The 
adoption of standard English names is a problem for any work covering such a wide area, 
many parts of which have been dealt with in a variety of past handbooks. In each case a 
preferred name has been adopted, others that have been used being cited in the 
annotations. 


Gillham, E. & B. 1996. Hybrid Ducks: a contribution towards an inventory. Pp. 88, 53 
colour and 3 black-and-white photographs. Published by the authors (P.O. Box 563, 
Wallington, Surrey SM6 9DX). ISBN 0-9511556-0-2. £14, plus packing and postage 
£1 (Europe), £2 (other continents). 21 x 15 cm. 


The outcome of a long-standing interest in hybrid ducks by the senior author, 
summarising data on 161 hybrid combinations, many of which are in the authors’ 
growing collection of live hybrid ducks. For each hybrid the plumage and other 
characters are described in detail, many being illustrated. There is a selective list of 225 
references. 


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Registered Charity No. 279583 


CONTENTS 


CLUB NOTICES.) 5 526 Sos ota OY Siete eo 
CLANCEY, P. A. Subspeciation in Layard’s Tit-babbler of the southwestern 
APfrotropies i. G2. as ieee RS a ere ore ae 
KRABBE, N., POULSEN, B. O., FR@LANDER, A. & BARAHONA, O. R. Range 
extensions of cloud forest birds from the high Andes of Ecuador: new 

sites for rare or little-recorded species .... ..5. 2.5...) oe 
TOYNE, E. P. & FLANAGAN, J. N. M. Observations on the breeding, diet and 
behaviour of the Red-faced Parrot Hapalopsittaca pyrrhops in southern 
Beeuador eo) eee 2 sls Sy an 0 5 gat erence toa ta 
SMITH, P. W. ‘The history and taxonomic status of the Hispaniolan Crossbill 
Loxia megaplaga oii... oe ee aw Oe vine oe eee 
VALLELY, A. C. & AVERSA, T. New and noteworthy bird records from Belize 
including the first record of Chestnut-collared Swift Cypseloides rutilus 
LOWEN, J. C., CLAY, R. P., BARNETT, J. M., MADRONO, N., A., PEARMAN, M., 
LANUS, B. L., TOBIAS, J. A., LILEY, D. C., BROOKS, T. M., ESQUIVEL, E. Z. & 
REID, J. M. New and noteworthy observations on Paraguayan avifauna 
RAPOSO, M. A. & PARRINI, R. On the validity of the Half-collared Sparrow 
Arremon semitorquatus Swainson, 1837... .)...... «. 2...) 
COWAN, P. J. What.1s.a desert bird? ....) 2 2...)5.. ss 24 See 
GREGORY, P. Range extensions and unusual sightings from Western 
Province, Papua New Guinea ..... 2.0.6.2. ..0.% 0 iJ 

In Brief REINO, L. M. & PATACHO, D. A new breeding area for the Song 
Thrush Turdus Philomelos in north Portugal ............. 

GONZALEZ M., O. First description of the nest of the Slender- 

billed Finch yo...) esa age ee aes ee oe 

DESFAYES, M. A specimen of Hirundo abyssinica from Madagascar 
BRADSHAW, C. G., KIRWAN, G. M. & WILLAMS, R. S. R. First record 

of Swainson’s Hawk Buteo swainsoni for the West Indies... . 

MORAES, V..\S. & KRUL, R. Notes on the Black-backed Tanager 

Tangara peruviana (Desmarest, 1806) .......... 2.5 sae 

BOOKS RECEIVED (10.6 406 a eis es we ede os ee wee hg Yeon 


248 


Zsa 


264 


272 


ZI3 


294 
299 


304 


312 


314 
315 


315 


316 
319 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 


mail, after their current subscription has been paid. 


COMMITTEE 
Revd T. W. Gladwin (Chairman) (1997) Miss H. Baker (1994) 
Mrs A. M. Moore (Vice-Chairman) (1997) Dr R. P. Pr¥s-Jones (1995) 
Cdr M. B. Casement, OBE, RN N. H. F. Stone (1995) 
(Hon. Secretary) (1996) D. Griffin (1997) 
D. J. Montier (Hon. Treasurer) (1997) N. J. Redman (1997) 


Hon. Editor: Prof C. J. Feare 
Chairman of Publications Sub-committee: Mrs A. M. Moore 
Publications Officer: S. J. Farnsworth 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


~~ 


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pSaids ee. of the 


BRITISH 


DR gots TECL Un 


EDITED BY 


Prof Chris Feare 


Volume 118 
1998 


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LIST OF AUTHORS AND CONTENTS 
AGRO, D.J. & RIDGELY, R.S. First record of the Striped Manakin 


Maskneropterus vesuins in Giryana® . 2.) 05. SPR. SOE oe 122 
ANDERSON, D.L., BONTA, M. & THORN, P. New and noteworthy bird 
eeEES IT OTA Ea ONCOUEAS 65 se! = ayshs, 5) s%- aos 5 WSR A OREN Ear oS DA DY 178 


ARAYA, T.A. see COLLINS, C.T. 
ARCHER, A.L. & ILES, D.B. New bird records for Unguja (Zanzibar Island). 166 
BEGAZO, A.J. & VALQUI, T. H. Birds of Pacaya-Samiria National Reserve 

with a new population (Myrmotherula longicauda) and new record for Peru 


SNM UPURRLEES SVIRECIRCTOUS ) ss 5 oes = 5 5 ss 2 os oe RN Be oes, SERIO LR 159 
BONTA, M. see ANDERSON, D.L. 
eee TUNER ED LGR. CFI TEL). BIN VAY DARREN MO Mad, Doe 62, 66, 200, 261 
BRACE, R.C. & HORNBUCKLE, J. Distributional records of and indentifi- 

cation notes on birds of the Beni Biological Station, Beni, Bolivia.......... 36 
CARTER, M. F. & DICKERMAN, R.W. The first documented nest and eggs of 

ems anselicr: mentalis... 001, AUR OO Ot? SE OR aL BE, 123 


CHRISTIDIS, L. see McALLEN, I.A.W. 
CLARK, R. see CLARK, W.S. 
CLARK, W.S. & CLARK, R. Differences in tarsal length between adult female 


Montagu’s and Pallid Harriers: an easy method to separate specimens...... 52 
CLARK, W.S & WHEELER, B.K. ‘Dark-morph’ Sharp-skinned Hawk reported 
from California is normal juvenile female of race perobscurus .............. 191 


CLAY, R.P. see LANUS, B.L. 
CLUB NOTICES 


Report of the Committee for 1997 & A.G.M. Agenda .................... 1 
Annual General Meeting 19 May 19989. 2.0. 2.220201 ie ee es. 129 
Financial Statement for the year ended 31 December 1997................ 137 
EE IPMEIOIEESS OP sw wa we nw cys ots ee RE ete Dey S98. Ad) 4, 65, 133, 201 
COLLAR, N.J. Wattled Cranes in Guinea-Bissau "7.5. 222 0) of). 57 
COLLAR, N.J. Monotypy of Francolinus griseostriatus ..........00 0c 124 
COLLINS, C.T. & ARAYA, T.A. Natal pterylosis of tanagers II: Tachyphonus, 
uerusiang Tangara ......... .. .FhackstAsten EE ee at eS, 172 


COLLINS, C.T. Food delivery and chick provisioning in the cypseloidine swifts. 108 
DELESTRADE, A. Distribution and status of the Ethiopian population of the 


Giese? yrenbcorax pyrrhocorax bailey? ‘eros Ver. Oe OG sso 101 
DEMEY, R. & FISHPOOL, L.D.C. On the existence of a melanistic morph of 
the Long-tailed Hawk Urotriorchis macrourus .........0.20. cece eenceces 105 


DIAS, R.A. see MAURICIO, G.N. 

DICKERMAN, R.W. see CARTER, M.F. 

ERRATA: Bull.Brit.Orn.Cl. 117(1) 1997 
Bull.Brit.Orn.Cl. 118(2) Plate 1. 
rere eter A 21RD) IGT 82. ee a an oe ces cly hae Nekedulyaet at (Po EEE 140 

FIGUEROA-ESQUIVEL, E.M., NAVARRO, S., A.G. & POZO-DE LA 
TIJERA, C. New distributional information on the birds of southern 
DEG Et. AICRIEO. .. 2). ne CI. ho fuccod cles oh tao ae. BA PR 32 

FISHPOOL, L.D.C. see DEMEY, R. 

FRITH, C.B. & FRITH, D.W. Hybridization between Macgregor’s Bowerbird 
Amblyornis macgregoriae and the Streaked Bowerbird A. subalaris (Ptilono- 
eae) OF Neg (suined |. o.oo oan chk aon KS CREO UEE OL Fan z, 

FRITH, D.W. see FRITH, C.B. 

GAMAUF, A. & PRELEUTHNER, M. A new taxon of the Barred 


Honeybuzzard Pernis celebensis from the Philippines....................-. 90 
GARNETT, S. Clarification of the type locality of the Golden-shouldered Parrot 

Oe ea aie en ee? <5 32) 6) ae Ob ee rr 196 
GREGORY, S.M.S. The correct citation of Coragyps (Cathartinae) and Ardeotis 

MN se no cm slichin Shige metal wise e ARM OE SIRE Ses Sew lag mnie e's gman 126 


HICKS, D. see RILEY, J. 

HOLMGREN, J. A. parsimonious phylogenetic tree for the swifts, Apodi, 
compared with DNA- ALEANY RAG UNG MS EIMLE NS cee ne nT A gh Sy oes sei p-iscm: vi: va th 238 

HORNBUCKLE, J. see BRACE, R.C. 


iv 


HORNE, J.F.M. & SHORT, L.L. A case of ‘song-capture’: Rufous-naped Lark 
uses nightjar song in place of its own song...!/..-5....-.-.=.>-\4 eee 

ILES, D.B. see ARCHER, A.L. 

JESSUP, L. ‘Gibsone’s nondescript’ and the lost type of Scissirosrum dubium 
(Latham) tractcctose bie -erost 4 -AGORT. 2.4 ATO ee 

KLUZA, D.A. see ZYSKOWSKI, K. 

LAMBERT, F.R. A new species of Amaurornis from the Talaud Islands, 
Indonesia, and a review of taxonomy of bush hens occuring from the 
Philippinesito, Australasia. 2. eugon? An. aba .l4. /D.. POLE eee 

LAMBERT, F.R. & RASMUSSEN, P.C. A new Scops Owl from Sangibe 
Island, Indonensia ... - o.oo see es ee eee oe ee ole eye eer 

LANUS, B.L., CLAY, R.P. & LOWEN, J.C. A new plumage of the 
White-winged Nightjar Caprimulgus candicans (Aves: Caprimulgidae) ...... 

LECROY, M. & PECKOVER, W.S. Misima’s missing birds ................. 

LOWEN, J.C. see LANUS, B.L. 

MAURICIO, G.N., & DIAS, R.A. Range extensions and new records for forest 
birds in southern Rio Grande do Sul, Brazil.............0.0... 00002000: 

McALLAN, I.A.W. & CHRISTIDIS, L. Neotype of the Hooded Plover 
Charadrius rubricollis Gmelin, 1789...............8.01. AGA DO eee 

NAVARRO, S., A.G. see FIGUEROA-ESQUIVEL, E.M. 

OLSON, S.L. Notes on the systematics of the Rockrunner Achaetops 
(Passeriformes, Timaliidae) and its presumed relatives. ................... 

OLSON, S.L. Lectotypification of Charadrius rubricollis Gmelin 1789.......... 

PACHECO, J.F. & WHITNEY, B.M. Correction of the specific name of 
Long-trained Nightjar... 5.2.20. 0. cp e ee nes el ee se 2 9 oe 

PECKOVER, W.S. see LECROY, M. 

PETERSON, A.T. The distribution and type locality of the extinct Slender-billed 
Grackle Quiscalus palustris schcnsuh? 14. baghce» tows stl. 2b 7a See 

PETERSON, A.T. see ZYSKOWSKI, K. 

POZO-DE LA TIJERA, C. see FIGUEROA-ESQUIVEL, E.M. 

PRELEUTHNER, M. see GAMAUF, A. 

RASMUSSEN, P.C. A new Scops-owl from Great Nicobar Island ............ 

RASMUSSEN, P.C. see LAMBERT, F.R. 

REID, K. Franklin’s Gull Larus pipixcan at south Georgia................++-- 

RIDGELY, R.S. see AGRO, D.J. 

RILEY, J.. HICKS, D. & WARDILL, J.C. The taxonomic status of Halcyon 
enigma on the Talaud Islands, Indonesia .................020 eee ee eeeee 

RILEY, J., HICKS, D. & WARDILL, J.C. The identity of Eos histrio challengeri 
Salvadori: 1890 vii. cies ee alae ee ee os 5 owe ne PN eee gre ene 

SHORT, L.L. see HORNE, J.F.M. 

STILES, F.G. Notes on the biology of two threatened species of Bangsia tanagers 
in northwestern, Colombia... . 5.6.24. ss + ot + age les ee 

THORN, P. see ANDERSON, D.L. 

TYRBERG, T. The date of publication of Montin’s description of Lagopus mutus. 

VALQUI, T.H. see BEGAZO, A.J. 

VAN BALENS Si‘A‘hy brid muna? 222900 PD). 0 ee 

WALTERS, M. The eggs of the Pink-headed Duck ......................... 

WARDILL, J.C. see RILEY, J. 

WELLER, A-A. On types of trochilids in the Natural History Museum, Tring I. 
Amazilia Sumichrasti Salvin, in relation to morphology and biogeography 
within the A. beryllina complex. ....<.........+-. GORI A Reh ae Aeon 

WHEELER, B.K. see CLARK, W.S. 

WHITNEY, B.M. see PACHECO, J.F. 

ZYSKOWSKI, K., PETERSON, A.T. & KLUZA, D.A. Courtship behaviour, 
vocalizations and species limits in Atthis hummingbirds .................-. 


153 


256 
259 


it9 


141 


55 


114 
193 


za 
56 


118 
187 


249 


INDEX TO SCIENTIFIC NAMES 
(Compiled by Mary N. Muller) 


All generic and specific names (of birds only) are indexed. Names of new taxa are indexed in 
bold print under generic, specific and, where appropriate, subspecific names. 


Accipiter collaris 27 

— gentilis 192 

— novaehollandiae 221, 231-2 

— ovampensis 191 

— poliocephalus 221, 232 

— striatus 191-3 

— superciliosus 40 

— tachiro 191 
Achaetops pycnopygius 47-51 
Acrocephalus gracilirostris 171 

— schoenobaenus 171 

— acuminata, Calidris 65, 221 

— acunhae, Nesospiza 134 
Aegithalos caudatus 202 
Aegotheles spp. 65 

— albertisi 66 

— insignis 66, PI.1. 
aenea, Chloroceryle 181 

— Glaucis 181 
Aerodramus sp. 239, 245 

— bartachi 246-7 

— brevirostris 246-7 

— elaphrus 246-7 

— francicus 246-7 

— fuciphagus 246-7 

— maximus 246-7 

— salanganus 246-7 

— sawtelli 246-7 

— spodiopygius 243, 246-7 

— terraereginae 246-7 
Aeronautes sp. 239, 243-4 
aeruginosus, Circus 54, 168 
afer, Sphenoeacus 48-9 
affinis, Apus 245-7 

— Aythya 189 

— Euphonia 182 
afra, Pytilia 172 
africana, Mirafra 153-8 
Ailuroedus spp. 7 
akool, Amaurornis 71, 73 
albertisi, Aegotheles 66 
albescens, Synallaxis 41 
albicollis, Xiphocolaptes 20 
albifrons, Donacospiza 45 

— Sterna 170 
albigula, Myzomela 223, 228, 232 
albigularis, Laterallus 180 
albinucha, Xenopsaris 44 
albolimbatus, Megalurus 66 
Alcedo atthis 223, 230-1 

— azurea 236 

— ispida 231 
alcyon, Ceryle 120 
alecto, Myiagra 223, 232, 235 
alfredi, Otus 148 
alius, Otus sp.nov. 143-51, 214 


Amalocichla sclateriana 66 
Amandava amandava 153 
Amaurornis spp. 67-80 

-— akool 71, 73 

— isabellinus 69-79 
Amaurornis magnirostris. sp.nov. 74- 

80, P1.2 

— moluccanus 67-80, P1.2 

— olivaceus 67-80, P1l.2, 221, 224 
ae spp. 89 

beryllina 249-55 

— cyanocephala 252 

— luciae 181 

— ocai 252 

— rutila 255 

— sumichrasti 249-55 

— violiceps 255 

— viridifrons 255 
amazonicus, Thamnophilus 163 
Amblyornis spp. 7-8 

— flavifrons 7 

— inornatus 7 

— macgregoriae 7-13, 66 

— subalaris 7-13 
amboinensis, Macropygia 222, 229 
americana, Aythya 189 
Amphilais seebohmi 49 
anaethetus, Sterna 222 
Anas spp. 190 

— hottentota 168 

— superciliosa 221, 229 
Anastomus lamelligerus 168 
andrei, Chaetura 18 
angolensis, Oryzoburus 33-4 
angustirostris, Marmaronetta 189 
Anisognathus spp. 27 
Anous sp. 222, 233 

— minutus 65 

— stolidus 65, 170 
Anthus ludovicianus 120 
apivorus, Pernis 97-8 
Aplonis cantoroides 223, 229, 235 

— metallica 223, 229, 232 
Dus sp. 239, 241, 243- 5 

affinis 245- i 

— apus 109, 170, 240 

- nipalensis 246-7 

— pallidus 135 

— unicolor 135 
arcae, Bangsia 30 
Archilochus spp. 83, 88 
Ardea melanocephala 167 
ardens, Selasphorus 88 
Ardeotis spp. 126 

— australis 66 
ariel, Fregata 167, 221 


armatus, Vanellus 134 
arthus, Tangara 174-5 
asiaticus, Ephippiorhynchus 66 
Asio clamator 180 
Athene blewitti 201 

— brama 202 


aspasia, Nectarinia 220, 223, 229, 232, 235 


Atlantisia rogersi 134 
atriceps, Hypergerus 135 
atricilla, Larus 55 
Atthis spp. 82-9 

-— ellioti 82-9 

— heloisa 82-9 

— morcomi 85 
atthis, Alcedo 223, 230, 231 
aurantius, Chaetops 48 
aureocincta, Bangsia 25-31 
aurulentus, Piculus 19 
australis, Ardeotis 66 

— Aythya 189 
Automolus ochrolaemus 33 
Aythya affinis 189 

americana 189 

— australis 198 

— baeri 189 

— collaris 189 

— ferina 189 

— fuligula 189 

— innotata 189 

— marila 189 

— novaeseelandiae 189 

— nyroca 189 

valisineria 189 

azurea, Alcedo 236 


baeri, Aythya 189 
bakkamoena, Otus 149 
balasiensis, Cypsiurus 246-7 
Balearica regulorum, 168 
Bangsia arcae 30 

— aureocincta 25-31 

— edwardsi 30 

— melanochlamys 25-31 
barbarus, Laniarius 135 
bartachi, Aerodramus 246-7 
beccarii, Otus 146-7, 213 
bergii, Sterna 222 
beryllina, Amazilia 249-55 
bicolor, Ducula 222, 229 
bitorquatus, Rhinoptilus 201 
blewitti, Athene 201 

— Heteroglaux 201 
borbonica, Phedina 242 
Bowdleria punctata 49 
brachyura, Myrmotherula 163 
brachyurus, Buteo 17 
Bradypterus luteoventris 49 
brama, Athene 202 
brasiliana, Hydropsalis 260 
brevipes, Tringa 221 
brevirostris, Aerodramus 246-7 
brunneus, Dromaeocercus 49 


Bugeranus carunculatus 57-8 
burmeisteri, Phyllomyias 21 
Buteo brachyurus 17 


cabanisi, Ceryle 120 
Cacatua galerita 222, 235 
caerulescens, Ptilorrhoa 65 

_ Sporophila 45 
Calidris acuminata 65, 221 

— ferruginea 221, 224 

— ruficollis 65, 221 
Caloenas nicobarica 222, 233 
Calypte spp. 83, 88-9 

— costae 85 
campanisona, Chamaeza 20 
Campephilus robustus 19 
canadensis, Sakesphorus 163 
candicans, Caprimulgus 183-6 
caniceps, Myiopagis 164 
canorus, Cuculus 170 
cantoroides, Aplonis 223, 229, 235 
Caprimulgus sp. 154 

— candicans 183-6 

— europaeus 170 

— forcipatus 259 

— fraenatus 157 

— parvulus 40 

— poliocephalus 154 
Capsiempis flaveola 164 
carbo, Ramphocelus 173-5 
Cardinalis cardinalis 182 
cardinalis, Myzomela 235 
Carpornis cucullatus 16, 22-3 
carunculatus, Bugeranus 57-8 


caryophyllacea, Rhodonessa 187-90, 201 


castanotis, Pteroglossus 16 
Casuarius casuarius 66 
caudatus, Aegithalos 202 
cauta, Diomedea 167 
cayana, Tangara 174-5 
cayanensis, Leptodon 179 
celebensis, Pernis 90-9 


celebensis winkleri, Pernis subsp.noy. 


94-9 

Cercotrichas podobe 48 
cerviniventris, Chlamydera 65 
Ceryle alcyon 120 

— cabanisi 120 
Chaetops aurantius 48 

— frenatus 48-50 

— pyconpygtus 48-51 
Chaetura sp. 239, 243, 245 

— andrei 18 

— pelagica 245 

— vauxi 246-7 
Chalcophaps indica 222, 229, 233 
chalybea, Euphonia 24 
Chamaeza campanisona 20 

— ruficauda 16 
Charadrius cucullatus 59-60, 256-9 

— dubius 169 

— rubricollis 59-60, 256-9 


Charmosyna papou 66 
Cheramoeca sp. 242 
chinensis, Oriolus 116 
Chlamydera spp. 7 

— cerviniventris 65 

— maculata 9 

— nuchalis 9 
Chloebia gouldiae 153 
chloris, Halcyon 113-8, 222, 229 

— Todiramphus 78 
Chloroceryle aenea 181 

— inda 181 
Chlorochrysa phoenicotis 27-8 
chloropus, Gallinula 134 
Chlorospingus flavigularis 28 

— semifuscus 27, 28 
Chlorothraupis stolzmannii 28 
Chrysococcyx lucidus 65, 222, 230 
chrysocephalus, Sericulus 9 
chrysomela, Monarcha 65 
chrysopterygius, Psephotus 196-9 
Ciccaba nigrolineata 180 
Cichladusa spp. 48 
Cicinnurus magnificus 66 

— regius 65 
Cinclus mexicanus 120 
cinerascens, Monarcha 223, 229 
Circus spp. 54 

— aeruginosus 54, 168 

— macrourus 52-4 

— pygargus 52-4 
cirrocephalus, Larus 55 
Cistothorus palustris 120 
clamator, Asio 180 
Cnemophilus macgregorii 66 
collari, Otus sp.nov. 204-16, Pl. 4 & 5 
collaris, Accipiter 27 

— Aythya 189 

— Sporophila 45 
Collocalia sp. 239, 244-5 

— esculenta 222, 231, 243, 246-7 

— linchi 246-7 

— troglodytes 246-7 

— vanikorensis 222, 229 
Colluricincla sanghirensis 204 
colma, Formicarius 15 
Colonia colonus 16 
colonus, Colonia 16 
Columba junoniae 134 

— palumbus 134 

— trocaz 134 

— vitiensis 222, 228 
columbarius, Falco 192 
comeri, Porphyriornis 134 
Conopias trivirgata 165 
Conopophaga lineata 20-1 

— peruviana 164 
contaminatus, Heliobletus 24 
Copsychus spp. 48 
Coracina novaehollandiae 223, 230 

— tenuirostris 223, 230 
Coragyps spp. 126 


coromanda, Halcyon 115 
coronatus, T'achyphonus 24 
Corvus orru 223 

— splendens 168 
costae, Calypte 85 
Cranioleuca gutturata 41 
Crax globulosa 161 

— mitu 162 
creagra, Hydropsalis 259-60 

— Macropsalis 259-60 
cristata, Galerida 157 
cucullatus, Carpornis 16, 22-3 

— Charadrius 59-60, 256-9 
Cuculus canorus 170 

— gularis 170 
cunicularia, Speotyto 40 
cuvieri, Falco 168 
cyanocephala, Amazilia 252 

— Psittacula 201 
cyanocephalus, Malurus 65 
cyanoleuca, Myiagra 223 

— Notiochelidon 177 
Cyanopica cyanus 135 
cyanus, Cyanopica 135 
cypriaca, Oenanthe 134 


Cypseloides spp. 109-11, 239, 245 


— niger 110, 246-7 

— rutilus 109 
Cypsiurus sp. 239, 243 

— balasiensis 246-7 


dactylatra, Sula 167 
danae, Tanysiptera 66 
Delichon sp. 242 
Dendrocolaptes platyrostris 20 
Dicaeum pectorale 223, 232 
dichrous, Pitohui 66 
dicolorus, Ramphastos 18-9 
Diomedea cauta 167 
dominicanus, Larus 55 
Donacospiza albifrons 45 
dougallii, Sterna 170 
Dromaeocercus brunneus 49 
Drymophila rubricollis 16 
Dryocopus lineatus 19 
dubium, Scissirostrum 5-6, Pl. 1 
dubius, Charadrius 169 
Ducula spp. 65 

— bicolor 222, 229 

— pinon 220, 222, 233 

— pistrinaria 222, 229 

— zoeae 218, 228, 234 
Dupetor flavicollis 221 
Dysithamnus mentalis 41 


edwardsi, Bangsia 30 

Egretta intermedia 221, 232 
— sacra 221 

Elaenia sp. 177 

Elanoides forficatus 33 

elaphrus, Aerodramus 246-7 

elegans, Otus 150, 213 


ellioti, Atthis 82-9 
enganensis, Otus 143, 145, 149, 211, 214 
enigma, Halcyon 113-7 

— 'Todiramphus 78 
Eos histrio 193-6 
Ephippiorhynchus asiaticus 66 
episcopus, Thraupis 173-7 
eremita, Megapodius 65 

— Nesocichla 134 
Erithacus luscinia 177 
erythrophthalma, Netta 189 
erythrorhynchos, Pelecanus 179 
Esacus magnirostris 220-1 
esculenta, Collocalia 222, 231, 243, 246-7 
Eudynamys taitensis 222, 229 
EBuphonie affinis 182 

chalybea 24 

— hirundinacea 182 

— minuta 182 

— violacea 24, 173-4 

— xanthogaster 28 
europaeus, Caprimulgus 170 
Eurystomus orientalis 223 


Falco columbarius 192 

— cuvieri 168 

— peregrinus 221, 233 
fasciata, Melospiza 120 
fasciatus, Phyllomyias 21 
ferina, Aythya 189 
ferruginea, Calidris 221, 224 
finschii, Psittacula 201 
fischeri, Vidua 172 
flammula, Selasphorus 88 
flaveola, Capsiempis 164 
flavicollis, Dupetor 221 
flavifrons, Amblyornis 7 
flavigularis, Chlorospingus 28 
Florisuga mellivora 33 
forcipata, Macropsalis 260 
forcipatus, Caprimulgus 259 
forficatus, Elanoides 33 
Formicarius colma 15 
fraenatus, Caprimulgus 157 
francicus, Aerodramus 246-7 
Francolinus griseostriatus 124-6 

— leucoscepus 154 
Fregata ariel 167, 221 

— minor 167 
frenatus, Chaetops 48-50 
freycinet, Megapodius 221, 229, 233 
frontalis, Synallaxis 41 
fuciphagus, Aerodramus 246-7 
fuertesi, Icterus 34 
fuliginosa, Psalidoprocne 242 
fuligula, Aythya 189 
fulva, Pluvialis 65, 169, 220-1, 233 
fulviventris, Hylopezus 181 
fuscata, Sterna 65, 170 


galatea, Tanysiptera 65 
Galerida cristata 157 


galerita, Cacatua 222, 235 
Gallinula chloropus 134 
Gampsonyx swainsonii 40 
gentilis, Accipiter 192 
geoffroyi, Geoffroyus 222, 229 
Geoffroyus geoffroyi 222, 229 
Geotrygon montana 18 
Gerygone magnirostris 223, 229, 230-1 
gigas, Hydrochous 241, 243, 246-7 
Glareola maldivarum 221, 224 
Glaucis aenea 181 
globulosa, Crax 161 
goughensis, Rowettia 134 
gouldiae, Chloebia 153 
Goura scheepmakeri 66 
gracilirostris, Acrocephalus 171 
griseostriatus, Francolinus 124-6 
griseotinctus, Zosterops 223, 227-8, 231 
Grus rubicunda 66 
guianensis, Morphnus 179 
gujanensis, Synallaxis 41 
gularis, Cuculus 170 
gunningi, Sheppardia 171 
guttula, Monarcha 223, 228-30, 232 
gutturata, Cranioleuca 41 
Gymnocrex spp. 67-9 

— talaudensis 78-80 


haematodus, Trichoglossus 65, 222, 229, 
234 
Haematopus longirostris 221 
ee chloris 113-7, 222, 229 
coromanda 115 

— enigma 113-7 

— macleayi 222, 230 

— sancta 65, 113, 222, 230 

— saurophaga 223, 230 
Haliaeetus leucogaster 221, 233 

— vocifer 167 
haliaetus, Pandion 120, 221 
Haliastur indus 221, 233 
hamatus, Rostrhamus 161 
Harpia harpyja 179 
harpyja, Harpia 179 
Harpyopsis novaeguineae 66 
Heliobletus contaminatus 24 
heloisa, Atthis 82-9 
Hemiprocne sp. 239, 244-5 

— mystacea 246-7 
Hemitriccus iohannis 164 
Heteroglaux sp. 202 

— blewitti 201 
himalayana, Psittacula 201 
Himantopus himantopus 221, 230 
Hippolais pallida 171 
Hirundapus sp. 239, 243, 245 
hirundinacea, Euphonia 182 
Hirundo sp. 242 

— nigricans 242 

— rustica 171 
histrio, Eos 193-6 
hottentota, Anas 168 


Hydrochous sp. 239, 243-5 

gigas 241, 243, 246-7 

Hydropsalis brasiliana 260 

creagra 259-60 

torquata 260 

Hylopezus fulviventris 181 

nattereri 16 

hylophila, Strix 18 

Hylophilus pectoralis 44 

semicinereus 159, 163, 165 

Hypergerus atriceps 135 

hypochroma, Sporophila 45 

hypoinochrous, Lorius 218, 220, 222, 232, 
234 

hypoleucos, Tringa 221, 229 

hypospodia, Synallaxis 41 

hypoxantha, Sporophila 45 


Icterus fuertesi 34 

spurius 34 

Ictinia mississippiensis 32 

inda, Chloroceryle 181 

indica, Chalcophaps 222, 229, 233 
indus, Haliastur 221, 233 

inepta, Megacrex 66 

Inezia inornata 43 

innotata, Aythya 189 

inornata, Inezia 43 

inornatus, Amblyornis 7 

insignis, Aegotheles 66, PI.1 
insularis, Otus 141, 145, 147, 213 
intermedia, Egretta 221, 232 
Psittacula 201 

iohannis, Hemitriccus 164 
isabellinus, Amaurornis 69-79 
ispida, Alcedo 231 


jacquacu, Penelope 162 
jardineii, Turdoides 49 
junoniae, Columba 134 


krameri, Psittacula 201 


Lagopus mutus 56-7 

Lalage leucomela 223 ; 
lamelligerus, Anastomus 168 
Laniarius barbarus 135 

Larus atricilla 55 

cirrocephalus 55 
dominicanus 55 
maculipennis 55 

pipixcan 55-6 

ridibundus 55, 169 
scoresbii 55 

serranus 55 

Laterallus albigularis 180 
ruber 180 

Leptodon cayanensis 179 
leucogaster, Haliaeetus, 221, 233 
leucomela, Lalage 223 
leucophrys, Zonotrichia 153 
leucoptera, Psophia 162 
Leucopternis semiplumbea 179 


leucopyga, Nyctiprogne 162 
leucoryphus, Platyrinchus 16 
leucoscepus, Francolinus 154 
leucotis, Melozone 182 

linchi, Collocalia 246-7 

lincolnii, Melospiza 153 

lineata, Conopophaga 20-1 
lineatus, Dryocopus 19 

Lioptilus spp. 50 

litsipsirupa, Turdus 134 

lobata, Tringa 257, 259 

lobatus, Lobipes 257-8 

Phalaropus 59 

Lobipes lobatus 257-8 

Lonchura pallidiventer 118-9 
longicauda, Myrmotherula 42, 159, 163 
longirostris, Haematopus 221 
Lorius hypoinochrous 218, 220, 222, 232, 234 
luciae, Amazilia 181 

lucidus, Chrysococcyx 65. 222. 230 
ludovicianus, Anthus 120 

lugubris, Myrmoborus 163 
Lurocalis semitorquatus 18 
luscinia, Erithacus 177 
luteoventris, Bradypterus 49 


macconnelli, Mionectes 42 
Macgregoria pulchra 66 
macgregoriae, Amblyornis 7-13, 66 
macgregorii, Cnemophilus 66 
Machaeropterus regulus 122 
macleayi, Halcyon 222, 230 
Macropsalis creagra 259-60 
forcipata 260 
Macropygia amboinensis 222, 229 
macrorrhina, Melidora 66 
macroura, Vidua 172 
macrourus, Circus 52-4 
Urotriorchis 105-7 
maculata, Chlamydera 9 
maculipennis, Larus 55 
magicus, Otus 141-7, 204-15, Pl.3 & 4 
magna, Sturnella 34, 120 
magnificus, Cicinnurus 66 
magnirostris, Amaurornis sp.nov. 74- 
80, P1.2 

Esacus 220-1 

Gerygone 223, 229-31 
Malacoptila rufa 163 
maldivarum, Glareola 221, 224 
Malurus cyanocephalus 65 
manadensis, Otus 143-50, 204-15, Pl. 4 
mantananensis, Otus 215 
marginalis, Porzana 169 
marila, Aythya 189 
Marmaronetta augustirostris 189 
masteri, Vireo 31 
maximus, Aerodramus 246-7 
Mearnsia sp. 239, 243 
Megacrex inepta 66 
Megalurus albolimbatus 66 
timoriensis 49 


Megapodius eremita 65 

— freycinet 221, 229, 233 
melanocephala, Ardea 167. 
melanochlamys, Bangsia 25-31 
Melanodera sp. 134 
melanoleucus, Spizastur 180 
melanothorax, Sylvia 134 
melanura, Pachycephala 228 
melanurus, Myrmoborus 163 
Melidora macrorrhina 66 
mellivora, Florisuga 33 
Melocichla mentalis 48-51 
Melospiza fasciata 120 

— lincoln 153 
Melozone leucotis 182 
menetriesii, Myrmotherula 163 
mentalis, Dysithamnus 41 

— Melocichla 48-51 

— Sphenoeacus 48 
mentawi, Otus 149, 214 
Merops muelleri 123 

— nubicus 171 

— ornatus 65, 223-4, 229-30 
metallica, Aplonis 223, 229, 232 
mexicana, Tangara 173-5 
mexicanus, Cinclus 120 

— Quiscalus 120-1 
Micrastur ruficollis 17 
Micropsitta pusio 222, 229-30 
minor, Fregata 167 
minuta, Euphonia 182 
minutus, Anous 65 

— Numenius 66 
Mhionectes spp. 42 

— macconnelli 42 

— oleagineus 42 
Mirafra africana 153-8 

— passerina 157 

— rufocinnamomea 154 
mississippiensis, Ictinia 32 
mitu, Crax 162 
Modulatrix spp. 50 


moluccanus, Amaurornis 67-80, Pl. 2 


Monarcha chrysomela 65 
— cinerascens 223, 229 
— guttula 223, 228-30, 232 


-— trivirgatus 223, 227, 229-30 


montana, Geotrygon 18 
morcomi, Atthis 85 
Morphnus guianensis 179 
muelleri, Merops 123-4 
munda, Serpophaga 43 
mutus, Lagopus 56-7 
Myadestes spp. 50 
Myiagra alecto 223, 232, 235 

— cyanoleuca 223 

— rubecula 223, 227 
Myiopagis caniceps 164 
Myrmoborus lugubris 163 

— melanurus 163 
Myrmotherula brachyura 163 

— longicauda 42, 159, 163 


— menetriesii 163 

— surinamensis 43, 163 

— unicolor 16 
mystacea, Hemiprocne 246-7 
mystaceus, Platyrinchus 20, 44 


Myzomela albigula 223, 228, 232 


— cardinalis 235 
— nigrita 223, 232 


nattereri, Hylopezus 16 
Neafrapus sp. 239 


Nectarinia aspasia 220, 223, 229; 232, 235 


Neocossyphus spp. 50 
Neopelma sulphureiventer 42 
Neopsiotis, Porphyriornis 134 
Nesocichla eremita 134 
Nesospiza acunhae 134 
— wilkinsi 134 
Netta erythrophthalma 189 

— peposaca 189 

— rufina 187, 189 
nubicus, Merops 171 ° 
nicaraguensis, Quiscalus 120 
nicobarica, Caloenas 222, 233 
niger, Cypseloides 110, 246-7 
nigricans, Hirundo 242 

— Sayornis 120 
nigrita, Myzomela 223, 232 
nigrolineata, Ciccaba 180 
nipalensis, Apus 246-7 
Notiochelidon cyanoleuca 177 


novaehollandia, Accipiter 221, 231-2 


— Coracina 223, 230 

— Scythrops 222, 235 
novaeseelandiae, Aythya 189 
novaeguineae, Harpyopsis 66 
nubicus, Merops 171 
nuchalis, Chlamydera 9 
Numenius minutus 66 

— phaeopus 218, 221 
Nyctiphrynus ocellatus 180 
Nyctiprogne leucopyga 162 
nyroca, Aythya 189 
Nystalus striolatus 40 


ocai, Amazilia 252 
ocellatus, Nyctiphrynus 180 
ochrolaemus, Automolus 33 
Oenanthe cypriaca 134 
oleagineus, Mionectes 42 
olivacea, Piranga 173-4 


olivaceus, Amaurornis 67-80, Pl. 2, 221, 224 


Ophrysia superciliosa 201 
orientalis, Eurystomus 223 
Oriolus chinensis, 116 


ornatus, Merops 65, 223-4, 229-30 


orru, Corvus 223 
Oryzoborus angolensis 33 
Otus alfredi 148 


Otus alius sp.nov. 143-51, 214, P1.3 


— bakkamoena 149 
beccarii 146-7, 213 


Otus collari sp.nov. 204-16, Pls. 4 & 5 
— elegans 150, 213 
— enganensis 143-9, 211, 214 
— insularis 141, 145-7, 213 
— magicus 141, 146-7, 204-15, Pl. 4 
— manadensis 143-50, 204-15, Pl. 4 
— mantananensis 215 
— mentawi 149, 214 
— scops 141 
— spilocephalus 148 
— sunia 141, 150 
— umbra 143-9, 
ovampensis, Accipiter 191 


Pachycephala melanura 228 

— pectoralis 223, 227-8, 231 
pallida, Hippolais 171 
pallidiventer, Lonchura 118 
pallidus, Apus 135 
palmarum, Thraupis 173-7 
palumbus, Columba 134 
palustris, Cistothorus 120 

— Quiscalus 119-21 
Pandion haliaetus 120, 221 
Panyptila sp. 239, 243-4 
papou, Charmosyna 66 
Parabuteo unicinctus 179 
Paradisaea raggiana 65 
paradisaea, Vidua 172 
parasiticus, Stercorarius 169 
parvulus, Caprimulgus 40 
passerina, Mirafra 157 

— Spizella 153 
pectorale, Dicaeum 223, 232 
pectoralis, Hylophilus 44 

— Pachycephala 223, 227-8, 231 
pelagica, Chaetura 245 
Pelecanus erythrorhynchos 179 
Penelope jacquacu 162 
peposaca, Netta 189 
peregrinus, Falco 221, 233 
Pernis apivorus 97-8 

— celebensis 90-9 


Pernis celebensis winkleri subsp.nov. 


94-9 

— ptilorhyneus 97 
peruviana, Conopophaga 164 
phaeopus, Numenius 221 
Phalaropus lobatus 59 
Phedina borbonica 242 
philippensis, Spizaetus 95 
Philomachus pugnax 169 
phoenicotis, Chlorochrysa 27-8 
Phyllomyias burmeisteri 21 

— fasciatus 21 

— virescens 22 
Phylloscopus sibilatrix 171 

— trochilus 171 
picta, Psittacella 66 
Piculus aurulentus 19 
pileatus, Piprites 16 
Pinarornis plumosus 48-50 


x1 


pinon, Ducula 220, 222, 233 
Pipile pipile 162 
pipixcan, Larus 55-6 
Piprites pileatus 16 
Piranga olivacea 173-4 
pistrinaria, Ducula 222, 229 
Pitohui dichrous 66 
platycercus, Selasphorus 86, 88 
Platyrinchus leucoryphus 16 

— mystaceus 20, 44 
platyrostris, Dendrocolaptes 20 
plumbea, Polioptila 165 
plumosus, Pinarornis 48-50 


_ Pluvialis fulva 65, 169, 220-1, 233 


— squatarola 218, 221, 229, 233 
podobe, Cercotrichas 48 
poliocephalus, Accipiter 221, 232 

— Caprimulgus 154 
Polioptila plumbea 165 
pomarinus, Stercorarius 169 
Porphyrio porphyrio 168, 221 
Porphyriornis comeri 134 

— nesiotis 134 
Porzana marginalis 169 

— spiloptera 134 
preciosa, Tangara 24 
Procnias tricarunculata 181 
Psalidoprocne fuliginosa 242 | 
Psephotus chrysopterygius 196-9 
Pseudocolopteryx sclateri 44 
Psittacella picta 66 
Psittacula cyanocephala 201 

— finschii 201 

— himalayana 201 

— intermedia 201 

— krameri 201 

— roseata 201 
Psophia leucoptera 162 
Pteroglossus castanotis 16 
Ptilinopus spp. 65 

— rivoli 222, 228-30 

— superbus 222, 229 
Ptilonorhynchus violaceus 9 
ptilorhyneus, Pernis 97 
Ptilorrhoa caerulescens 65 
pugnax, Philomachus 169 
pulchra, Macgregoria 66 
punctata, Bowdleria 49 
pusilla, Spizella 153 
pusio, Micropsitta 222, 229-30 
pycnopygius, Achaetops 47-51 

— Chaetops 48-51 

— Sphenoeacus 47-51 
pygargus, Circus 52-4 
Pyroderus scutatus 23 
Pyrrhocoma ruficeps 23 
Pyrrhocorax pyrrhocorax 101-4 
Pytilia afra 172 


Quiscalus mexicanus 120-1 
— nicaraguensis 120 
— palustris 119-21 


raggiana, Paradisaea 65 
Rallina tricolor 236 
Ramphastos dicolorus 18-9 
Ramphocelus carbo 173-5 
regius, Cicinnurus 65 
regulorum, Balearica 168 
regulus, Machaeropterus 122 
Rhaphidura sp. 239, 245 
Rhipidura rufifrons 223, 229 

— rufiventris 223, 229, 232 
Rhinoptilus bitorquatus 201 
Rhodonessa caryophyllacea 187-90, 201 
ridibundus, Larus 55, 169 
Riparia 242 

— riparia 171 
rivoli, Ptilinopus 222, 228-30 
robustus, Campephilus 19 
rogersi, Atlantisia 134 
roseata, Psittacula 201 
Rostrhamus hamatus 161 
Rowettia goughensis 134 
rubecula, Myiagra 223, 227 
ruber, Laterallus 180 
rubicunda, Grus 66 
rubricollis, Charadrius 59-60, 256-9 

— Drymophila 16 
rufina, Netta 187, 189 
rufa, Malacoptila 163 
ruficapilla, Synallaxis 20 
ruficauda, Chamaeza 16 
ruficeps, Pyrrhocoma 23 
ruficollis, Calidris 65, 221 

— Milcrastur 17 

— Sporophila 45 
rufifrons, Rhipidura 223, 229 
rufiventris, Rhipidura 223, 229, 232 
rufocinnamomea, Mirafra 154 
rufus, Selasphorus 88 

— 'Tachyphonus 173-5 
rustica, Hirundo 171 
rutila, Amazilia 255 
rutilus, Cypseloides 109 


sacra, Egretta 221 

Sakesphorus canadensis 163 
salanganus, Aerodramus 246-7 
sancta, Halcyon 65, 113, 223, 230 
sanghirensis, Colluricincla 204 
sasin, Selasphorus 88 

saundersi, Sterna 170 
saurophaga, Halcyon 223, 230 
sawtelli, Aerodramus 246-7 
Saxicola spp. 48 

sayaca, Thraupis 174 

Sayornis nigricans 120 
scheepmakeri, Goura 66 
Schiffornis virescens 22 
schoenobaenus, Acrocephalus 171 
Schoutedenapus 239, 241, 243 
scintilla, Selasphorus 88 
Scissirostrum dubium 5-6. Pl. 1 
sclateri, Pseudocolopteryx 44 


sclateriana, Amalocichla 66 
scops, Otus 141 
scoresbi, Larus 55 
scutatus, Pyroderus 23 
Scytalopus speluncae 21 
Scythrops novaehollandiae 222, 235 
seebohmi, Amphilais 49 
Selasphorus spp. 83, 88 

— ardens 88 

— flammula 88 

— platycercus 86, 88 

— rufus 88 

— sasin 88 

- scintilla 88 
semicinereus, Hylophilus 159, 163, 165 
semicollaris, Streptoprocne 110 
semifuscus, Chlorospingus 27, 28 
semiplumea, Leucopternis 179 
semitorquatus, Lurocalis 18 
Sericulus spp. 7 

— chrysocephalus 9 
Serpophaga spp. 43 

— munda 43 

— subcristata 43 
serranus, Larus 55 
Sheppardia gunningi 171 
sibilatrix, Phylloscopus 171 
Sicalis sp. 134 
Sitta whiteheadi 134 
speluncae, Scytalopus 21 
Speotyto cunicularia 40 
Sphenoeacus afer 48-9 

— mentalis 48 

— pycnopygius 47-51 
spilocephalus, Otus 148 
spiloptera, Porzana 134 
spinosus, Vanellus 134 
Spizaetus philippensis 95 
Spizastur melanoleucus 180 
Spizella passerina 153 

— pusilla 153 
splendens, Corvus 168 
spodiopygius, Aerodramus 243, 246-7 
Sporophila spp. 45 

— caerulescens 45 

— collaris 45 

— hypochroma 45 

— hypoxantha 45 

— ruficollis 45 
spurius, Icterus 34 
squatarola, Pluvialis 218, 221, 229, 233 
stagnatilis, Tringa 169 
Steganopus tricolor 258 
Stellula sp. 83, 88 
Stercorarius parasiticus 169 

— pomarinus 169 
Sterna albifrons 170 

— anaethetus 222 

— bergii 222 

— dougallii 170 

— fuscata 65, 170 

— saundersi 170 


— sumatrana 218, 222, 233 
Stizorhina spp. 50 
stolidus, Anous 65, 170 
stolzmannii, Chlorothraupis 28 
Streptoprocne spp. 109-11, 239, 245 

— semicollaris 110 

— zonaris 110, 246-7 
striatus, Accipiter 191-3 
striolatus, Nystalus 40 
Strix hylophila 18 
Sturnella magna 34, 120 
subalaris, Amblyornis 7-13 
subcristata, Serpophaga 43 
Suiriri suiriri 42 
Sula dactylatra 167 
sulphureiventer, Neopelma 42 
sulphurescens, Tolmomyias 22 
sumatrana, Sterna 218, 222, 233 
sumichrasti, Amazilia 249-55 
sunia, Otus 141, 150 
superbus, Ptilinopus 222, 229 
superciliosa, Anas 221, 229 

— Ophrysia 201 
superciliosus, Accipiter 40 
surinamensis, Myrmotherula 42, 163 
swainsonii, Gampsonyx 40 
Sylvia melanothorax 134 
Synallaxis albescens 41 

— frontalis 41 

— gujanensis 41 

— hypospodia 41 

— ruficapilla 20 


tachiro, Accipiter 191 
Tachornis sp. 239, 243-5 
Tachymarptis sp. 239, 243 
Tachyphonus coronatus 24 

— rufus 173-5 
taitensis, Eudynamys 222, 229 
talaudensis, Gymnocrex 78 , 80 
Tangara spp. 27-8 

— arthus 174-5 

_— cayana 174-5 

— mexicana 173-5 

— preciosa 24 
Tanysiptera danae 66 

— galatea 65 
Telecanthura 239 

— ussheri 171 
tenuirostris, Coracina 223, 230 
terraereginae, Aerodramus 246-7 
Tersina viridis 173-4 
Thamnophilus amazonicus 163 
Thraupis episcopus 173-7 

— palmarum 173-7 

— sayaca 174 
timoriensis, Megalurus 49 
Todiramphus chloris 78 

— enigma 78 
Tolmomyias sulphurescens 22 
torquata, Hydropsalis 260 
tricarunculata, Procnais 181 


Trichoglossus haematodus 65, 222, 229, 
234 


tricolor, Rallina 236 

— Steganopus 258 
Tringa brevipes 221 

— hypoleucos 221, 229 

— lobata 257-9 

— stagnatilis 169 
trivirgata, Conopias 165 
trivirgatus, Monarcha 223, 227, 229-30 
trocaz, Columba 134 
trochilus, Phylloscopus 171 
troglodytes, Collocalia 246-7 
Turdoides spp. 50-1 

— jardineii 49 
Turdus litsipsirupa 134 
Tyranneutes virescens 122 


umbra, Otus 143-9 
undulatus, Zebrilus 161 
unicinctus, Parabuteo 179 
unicolor, Apus 135 

— Myrmotherula 16 
Urotriorchis macrourus 105-7 
ussheri, Telecanthura 171 


valisineria, Aythya 189 
Vanellus armatus 134 

— spinosus 134 
vanikorensis, Collocalia 222, 229 
vauxi, Chaetura 246-7 
Vidua fischeri 172 

— macroura 172 

— paradisaea 172 
violacea, Euphonia 24, 173-4 
violaceus, Ptilonorhynchus 9 
violiceps, Amazilia 255 
Vireo masteri 31 
virescens, Phyllomyias 22 

— Schiffornis 22 

— Tyranneutes 122 
viridifrons, Amazilia 255 
viridis, Tersina 173-4 
vitiensis, Columba 222, 228 
vocifer, Haliaeetus 167 


whiteheadi, Sitta 134 

wilkinsi, Nesospiza 134 

winkleri, Pernis celebensis subsp.nov. 
94-9 


xanthogaster, Euphonia 28 
Xenopsaris albinucha 44 
Xiphocolaptes albicollis 20 


Zebrilus undulatus 161 

zoeae, Ducula 218, 228, 234 

zonaris, Streptoprocne 110, 246-7 
Zonotrichia leucophrys 153 

Zoonavena 239 

Zosterops griseotinctus 223, 227-8, 231 


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XIV 
CORRECTIONS TO TEXT 


May 1998 not May 1988 
xanthogaster not xanthogastra 
Oryzoborus funereus = Oryzoborus angolensis funereus 
Serpophaga not Serophaga 
inornata not ornata 
Passeriformes not Passiformes 
Tanysiptera not Tanysiptra 
Ptilorrhoa not Ptillorrhoa 
albertist not albertsi 
litsipsirupa not litsitsirupa 
jacquacu not jaquacu 
menetriesi not minitriesit 
pugnax not stagnatilis 
Pytilia not Pytelia 
cayanensis not cayenensis 
luciae not lucia 
1898a not 1989a 
haliaetus not haliacetus 
freycinet not freycinete 
Eudynamys not Eudynamis 


Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


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B%63 ISSN 0007-1595 
Bulletin of the 


British Ornithologists’ Club 


Edited by 
Prof C. J. FEARE and Dr D. W. SNOW 


SNT HSON AY 


APR Q & 1998 
LIBRARIES 


—— ae a, 


Volume 118 No. 1 March 1998 


FORTHCOMING MEETINGS 


Tuesday 21 April 1998. Dr W. R. P. (Bill) Bourne will speak on “Birds and Islands’. At 
the age of ten, Bill was evacuated to Bermuda, in the central western North Atlantic, for three 
years during World War II, where he sailed and fished and chased the local birds. Since then, 
in the course of a complicated career ending with an Honorary Research Fellowship at 
Aberdeen University, he has periodically managed to spend shorter periods watching both the 
land and seabirds of a variety of other islands, ranging from Bear Island and Cyprus, via the 
Cape Verde Islands and Ascension, to South Georgia and Juan Fernandez. He will summarise 
these experiences and draw some conclusions. 

Applications to the Hon. Secretary by 7 April, please. 


Tuesday 19 May 1998. ANNUAL GENERAL MEETING at 6 p.m., followed by a Club 
Social Evening. There will be no speaker, but Members are invited to bring along one or 
two slides (or a specimen!) of a bird of topical interest, and to speak for not more than 
5-10 minutes about it. The aim will be to generate discussion, and to facilitate the exchange of 
information between Members. 

Applications to the Hon. Secretary by 5 May, please, including subjects to be raised and any 
special facilities required. 


Tuesday 14 July 1998. R. E. (Bob) Scott will speak on “Bird reminiscences (in a lighter 
vein)’’. Bob started an interest in birds when he was introduced to a tame Jackdaw during 
war-time evacuation from London. As a school-boy in the early 1950s, egg-collecting first 
introduced him to the subject in detail, and he became closely involved with the London 
Natural History Society, serving for periods as junior committee member for their 
ornithological branch, and subsequently as recorder for south of the Thames. After a few years 
in the Zoology Department of the British Museum (Natural History), he took a post as warden 
at the RSPB Dungeness reserve. He has recently retired after serving 37 years with the RSPB, 
ending his career as Head of Reserves Management. During his time at Dungeness he served 
on the BTO’s Ringing and Migration Committee, and still retains an interest in the area, as a 
Trustee of the Dungeness Bird Observatory. He is currently Vice-President of the BOU. Since 
retirement, Bob has increased his foreign travel, now leading parties of bird-watchers to many 
different localities. Together with his wife Ann, he has a full lecturing programme throughout 
the country. 

Applications to the Hon. Secretary by 30 Fune, please. 


Future dates for 1998. Further meetings have been arranged for the following Tuesdays: 
15 September—Dr Nigel Collar on “Two Indian Enigmas’’, 13 October—Dr Ben Hatchwell 
on “The cooperative behaviour of Long-tailed Tits’’, and 17 November—Dr Bill Porteous on 
“Birds of the Humboldt Current’’. 


Meetings are held in the Sherfield Building of Imperial College, South Kensington, London 
SW7. The nearest Tube station is at South Kensington, and car parking facilities are available; a 
map of the area will be sent to members, on request. The cash bar is open from 6.15 p.m., and 
a buffet supper, of two courses followed by coffee, is served at 7.00 p.m. (A vegetarian menu 
can be arranged if ordered at the time of booking.) Informal talks are given on completion, at 
about 8.00 p.m. 


Overseas Members visiting Britain are particularly welcome at meetings. For 
details in advance, please contact the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1998 


Apart from single copies made for the purposes of research or private study, or criticism or 
review, as permitted under UK law, no part of this publication may be reproduced, stored 
or transmitted in any form or by any means, except with prior permission in writing of the 
publishers, or in accordance with the terms of licences issued by the Copyright Licensing 
Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; for 
address see inside back cover. 


1 Bull. B.O.C. 1998 118(1) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 118 No. 1 Published: 27 March 1998 


REPORT OF THE COMMITTEE FOR 1997 


Meetings. Eight evening meetings were held in 1997, in the 
Sherfield Building at Imperial College, London. A total of 297 (218 
members and 79 guests) attended these meetings, which represented an 
average attendance of 37. The programme of speakers during the year 
again covered a wide variety of ornithological subjects, and was 
presented by speakers both from Britain and from overseas. ‘The May 
meeting following the AGM was in the form of a social evening, during 
which informal short talks and brief discussions were contributed by 
twelve participants, on a wide range of topical subjects. 


Committee. The Committee met six times during the year, and the 
attendance was 73%. The chief topic for discussions concerned the 
progress of Club publications, and the activities of the Publications 
Sub-Committee (see below). Following the announcement by Imperial 
College of a steep increase in charges for room hire and catering, 
alternative meeting arrangements were actively considered, but an 
acceptable agreement was negotiated to continue at Imperial College, at 
least for 1998. Dinner charges were, however, increased accordingly. 
Following the resignation of Mr S. J. Farnsworth as Hon. Treasurer, 
the administration of Membership records and subscription payments 
was transferred to the Hon. Secretary, with the assistance of Miss H. 
Baker, as membership secretary. Mr D. J. Montier was elected Hon. 
Treasurer, at the AGM, and assumed responsibility for all accounting 
matters. Mr N. H. F. Stone has given valuable advice and help as Club 
archivist. 

Mr Peter Oliver, representing the Trustees of the Herbert Stevens 
Trust Fund met with the Committee to give advice on the performance 
of the Fund. The Committee is most grateful to Mr Nigel Crocker, Mr 
Richard Price and Mr Peter Oliver for their time and expertise in this 
matter. 

It is with great regret that the Committee reports the deaths in the 
past year of H. P. H. Johnson (1957-97) and G. R. Cunningham- 
van-Someren (1946—Hon. Life Member), also of J. G. Williams 
(1951-97), in January 1998. 


Publications Sub-committee. Two titles were published in 1997. 
Occasional Publications No. 2 Manuscripts and Drawings in the 
Ornithology and Rothschild Libraries of the Natural History Museum, 
completed in 1996, appeared early in January; and Occasional 
Publications No. 3 Avian Egg-shells: an Atlas of Scanning Electron 
Micrographs was published in mid-November. 

Sales of both titles, and of Occasional Publications No. 1 Extinct and 
Endangered Birds in the Collections of the Natural History Museum, 


Report of the Committee for 1997 2 Bull. B.O.C. 1998 118(1) 


Avian Systematics and Taxonomy, and Avian Taxonomy from Linnaeus 
to DNA (papers presented at the joint B.O.C. and Linnean Society of 
London meeting in March 1996, published as Volume 117(2) Bull. 
B.O.C.) have continued steadily throughout the year. 

Work continues on the possible Bird Atlas of Uganda, and a number 
of other publications are being actively explored. 

‘The Sub-committee was strengthened by the agreement of Professor 
Chris Feare and Nigel Redman to join it in September, and by the 
appointment of a Publications Officer, John Farnsworth, who has 
assumed responsibility for the non-Member Subscribers, sales of 
publications and Bulletin back-numbers, and other sales items. 

Because of other commitments, Dr David Snow resigned from the 
Sub-committee, when he relinquished the Editorship of the Bulletin. 
Dr Robert Prys-Jones agreed to take over the chairmanship from 
Amberley Moore, as from May 1988. 


Membership. There were 569 paid-up Members at 31 December 
1997—320 with addresses in the U.K., and 249 overseas (45 countries). 
Active recruitment from the newly-elected Members of the Union, and 
others attending dinner meetings as guests, resulted in 29 new 
Members joining, or re-joining the Club. 

‘The Committee decided that, in the interests of economy, the annual 
publication of Members’ Address Lists would be discontinued. The 
Hon. Secretary would prepare this list annually, and duplicated 
copies would be available for any Member requesting a copy, on 
payment of £1, to cover the costs of preparation and postage, but 
the complete list would be published, at four yearly intervals, with 
the Bulletin. 


The Bulletin. The 320 pages of Volume 117 contained 39 main 
papers and 13 “In Brief’ articles. ‘The June issue was devoted to a 
series of papers presented at a joint meeting of the Club and the 
Linnean Society, under the heading Avian taxonomy from Linnaeus to 
DNA, and contained major reviews of taxonomies, species concepts, 
and their implications for conservation. One new species, a vanga from 
Madagascar, was described and illustrated in colour, and details of a 
subspecies of Phylloscopus warbler, now elevated to full specific status on 
the basis of song, and responses to playback of songs of near relatives. 
While two new subspecies were described, the validity of others was 
questioned. Of 17 papers presenting new distributional data, 12 were 
from the neotropics. New information on the biology and behaviour of 
eight species was described, four of these were from the neotropics, 
including the first description of the nest of the Slender-billed Finch. An 
encouraging discovery of the apparently thriving population of Jerdon’s 
Bushchats on islands in the Mekong Delta was reported, and another 
paper refined our definition of “desert birds’’. The subjects of papers 
again reflected the global cover of Club interests, and the authorship of 
papers extended to 25 countries, from five continents. 

We are again grateful to Mary Muller who continues to compile the 
index, and to Michael Casement for preparing the Club Notes and 
Notices that preface each issue. 


Report of the Committee for 1997 3 Bull. B.O.C. 1998 118(1) 


Bulletin Sales. Sales to Non-member Subscribers totalled 142, 
with 18 in the U.K. and 124 overseas (31 countries). Of the latter, 51 
were to the United States of America, and 14 to Germany. 


Finance. The increase in subscription rates for both Members and 
non-Member Subscribers, which took effect on 1 January 1997, 
appears to have caused very little fall-out in overall membership, and 
will provide a valuable boost to the Club’s income. Production costs of 
the Bulletin continue to increase, especially with the introduction of 
colour plates from time to time 

Completion of two new publications in the same year inevitably had 
a significant impact on cash flow. Occasional Publications No. 2, 
published in January, and No. 3 in December required a combined 
outlay of almost £8,000, already partially offset by sales achieved at the 
year’s end. 

Accounts for 1997 are not yet available, but will be tabled at the 
AGM and published in the Bulletin. 


ANNUAL GENERAL MEETING 


The Annual General Meeting of the British Ornithologist’s Club will 
be held in the Ante-room of the Sherfield Building, Imperial College, 
London SW7 at 6 pm on Tuesday 19 May 1988. 


AGENDA 
1. Minutes of the 1997 Annual General Meeting (see Bull. Brit. Orn. 
Cl. 117(3): 153-156). 
Chairman’s report. 
Report of the Committee and Accounts for 1997. 
The Bulletin. 
The election of Officers. The Committee proposes that: 
(i) Mr D. J. Montier be re-elected as Honorary Treasurer, 

(11) Commander M. B. Casement, OBE, RN, be re-elected Hon. 
Secretary. 

(ii) Mr R. E. Scott be elected a member of the Committee vice 
Miss Helen Baker who retires by rotation, and is ineligible for 
re-election. 

6. Any other business of which notice shall have been given in 

accordance with Rule (12). 


Wn -& & bo 


By Order of the Committee 
MICHAEL CasEMENT, Honorary Secretary 


The eight hundred and seventy second meeting of the Club was held on Tuesday, 4 
November 1997 at 6.15 pm. 30 Members and 9 guests attended. 

Members present were: The Rev. T. W. GLapwIin (Chairman), Miss H. Baker, Sir 
David BANNERMAN Bt., Captain Sir Thomas BarRLow Br. R.N., J. W. BARRINGTON, 
Captain M. K. Barritt R.N., P. J. BELMaAN, I. R. Bishop, D. M. BrapLey, Cdr M. B. 
CASEMENT R.N., Professor R..J. CHANDLER, S. J. FARNSWORTH, J. E. Francis, C. A. R. 
Heim, G. P. Jackson, J. A. Jopiinc, I. T. Lewis, B. A. E. Marr (Speaker), D. J. 
Montier, Mrs A. M. Moores, R. G. Morcan, P. J. OLiver, C. M. C. Patrick, Dr W. G. 


Club Notes 4 Bull. B.O.C. 1998 118(1) 


Porteous, Dr R. P. Prys-Jongss, P. S. Repman, R. C. SELF, P. J. SELLar, N. H. F. STONE 
and S. J. R. RuMsEy. 

Guests attending were: Mrs G. Bonuam, Mrs C. R. CasEMENT, Mrs J. M. GLapwin, 
Lady BANNERMAN, Ms C. Horr, Mrs S. Lewis, A. M. Martin, Mrs M. Montier and 
P. J. Moore. 


BOC Meeting—4 November 1997 


After dinner, Tony Marr gave a presentation, illustrated with colour 
slides, entitled Atlantic Seabird Odyssey. He gave his talk the sub-title 
The Agony and the Ecstasy to suggest the difficulties of getting out to 
sea to study seabirds, and the pleasure derived from actually locating 
and watching them. 

Seabirds face many threats—pollution, fishing, predation, and 
habitat destruction and degradation. He illustrated particular examples 
of problems caused by pollution and fishing. There is still much to 
learn about seabird migration and lifestyles and excellent identification 
books are available, particularly Harrison’s two major works. 

‘There are two basic requirements for a successful pelagic trip—a 
suitable boat, and suitable bait. After much trial and error, he had 
found that game-fishing boats offered speed and manoeuvrability, while 
yachts had the advantage of silence. The best bait (or ‘chum’, as it was 
inexplicably named) consisted of an evil-smelling mixture of fish oil 
and fish pieces, mashed into a pulp, and hung in a net over the back of 
the boat to trail in the water. ‘The smell quickly attracts seabirds from 
miles around. 

‘The main part of the talk then showed the seabirds that he had found 
in different parts of the Atlantic. Around Madeira and the Canaries he 
had watched Fea’s (Soft-plumaged) Petrels, Bulwer’s Petrels, White- 
faced and Madeiran Storm Petrels, and Cory’s and Little Shearwaters, 
concentrating on the Salvage and Desertas Islands where landings in the 
dark had been undertaken. Off Dakar in Senegal, the westernmost 
point of Africa, recent autumn studies had revealed a large southwards 
passage of seabirds, which included thousands of Pomarine Skuas, hun- 
dreds of Long-tailed Skuas and Sabine’s Gulls, Cory’s and Cape Verde 
Shearwaters, Wilson’s and Bulwer’s Petrels and South Polar Skuas. 

His talk concluded with evocative slides of Antarctica and some of its 
wildlife. The journey between South America and the Falklands 
offered great opportunities to see albatrosses, including Wandering, 
Black-browed, Grey-headed and Light-mantled Sooty. On _ the 
Antarctic peninsula were breeding Wilson’s Petrels, Giant Petrels, 
South Polar Skuas and innumerable penguins. He ended with close-up 
pictures of Hump-backed Whales amidst the dramatic Antarctic 
scenery; a sight he described as both moving and humbling. Man was 
an alien in this unforgiving habitat. 


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Figure 1. Watercolour illustration of Scissirostrum dubium by George Gibsone, labelled: 
‘“Non-descript from the collection of M. Collone”’. The original was burnt at Thompson 
the bird stuffers. 


L. Fessop 5 Bull. B.O.C. 1998 118(1) 


‘Gibsone’s nondescript’ and the lost type of 
Scissirostrum dubium (Latham) 


by L. Jessop 
Received 12 December 1996 


Trivial marginalia sometimes provide information that is interesting to 
the historian in a way that was never intended by the original author. 
Such is the case of ‘Gibsone’s nondescript’ and Scissirostrum dubium. A 
marginal comment beside an unpublished watercolour painting, by an 
obscure provincial English artist, provided clues that led to the 
unravelling of at least part of the history of the holotype of a species. 

George Gibsone (c. 1762-1846) was an amateur, 'yneside-based artist 
who is best known for his many watercolours of mollusc shells (Jessop 
1996), but he also produced a single volume of watercolours of birds that 
is owned by the Natural History Society of Northumbria, and held in 
their library at The Hancock Museum. ‘The paintings are undated, but 
some are on paper that is watermarked with the date 1794. The front 
flyleaf contains some information in manuscript about the history of the 
volume, ““C.M. Adamson from his father, June 1849. Drawings of birds 
by Mr George Gibsone late of Newcastle’, and “‘Pres. by Miss C. 
Adamson 1937’’: 1.e. the volume was owned by members of the Adamson 
family until it was presented to the Natural History Society in 1937. 

One of the birds is labelled in manuscript ‘“‘Non-descript from the 
collection of M. Collone. The original was burnt at Thompson the bird 
stuffers’, and this bird may be called ‘Gibsone’s nondescript’. 

M. Collone is undoubtedly Charles Alexandre de Calonne 
(1743-1802), who fled to England from France in 1797. George 
Humphrey catalogued his collection in London (Anon [Humphrey] 
1797) before it was dispersed, the most valuable items being sold 
privately in 1801 and the remainder by auction in May, June and July 
of that year (Whitehead 1969, Chalmers-Hunt 1976). 

Among the many birds listed in Humphrey’s catalogue, there is one 
that may be the bird figured by Gibsone as a nondescript. It is: ““40. A 
bird probably of a new genus—Oiseau probablement d’une genre neuf”’. 

‘“Gibsone’s nondescript’ was referred to the Bird Group at the 
Natural History Museum at Tring for identification, and following a 
suggestion by Mr David Gibbs it was confidently identified as 
Scissirostrum dubium (Latham, 1802), a species of Sturnidae from the 
island of Sulawesi. 

Scissirostrum dubium was first described by Latham in his General 
Synopsis (Latham 1801) as the Dubious Shrike, from a specimen 
seen at “Mr Thompson’s in Little Saint Martin’s Lane, London but 
without any history of its manner or country annexed’’. In the same 
year, Latham bestowed the latin binomen Lanius dubius upon the 
Dubious Shrike in the supplement to his Index Ornithologus (Latham 


1802). The subsequent history of its taxonomy can be traced via Sharpe 
(1890). 


L. Fessop 6 Bull. B.O.C. 1998 118(1) 


‘There is a strong and obvious coincidence between Thompson the bird 
stuffer mentioned by Gibsone and Mr Thompson’s in Little Saint 
Martin’s Lane, London where Latham saw the bird on which he based 
his new species. The suspicion that the two Thompsons are in fact 
identical is strengthened by reference to Holden’s Triennial Directory 
for 1805-07, where John Thompson a ‘Natural History Preserver’ is 
listed at 5 Little Saint Martin’s Lane. There was, incidentally, also an 
auctioneer called ‘Thompson next door at number 3, who may or may 
not have been the same John Thompson. 

Setting aside the extremely remote possibility that Thompson had 
two specimens of Scissirostrum dubium, one described by Latham and 
another figured by Gibsone, it is reasonable to conclude that both men 
saw the same bird. 

The manuscript annotation to Gibsone’s painting is important 
because there is no information from the published literature about the 
prior history of the holotype of Scisszrostrum dubium or of its fate. We 
know from Gibsone that the bird originated in the collection of de 
Calonne and that it was destroyed while at ‘Thompsons. 

Although the holotype of Lanius dubius Latham, 1802 is now known to be 
no longer in existence, it is a distinctive species, and there is no taxonomic 
confusion surrounding its identity: designation of a neotype is, therefore, 
unnecessary. There appears to be no discrepancy between the current 
interpretation of Scisszrostrum dubium and Gibsone’s painting of the holo- 
type, and no taxonomic changes arise from the discovery of the painting. 


References: 

Anon [Humphrey, G.], 1797. Museum Calonnianum. Specification of the various articles 
which compose the magnificent museum of Natural History collected by M. de Calonne in 
France and lately his property. London. 

Chalmers-Hunt, J. M. 1976. Natural History Auctions 1700-1972. A register of Sales in 
the British Isles. London. 

Jessop, L. 1996. George Gibsone and his conches. Trans. Nat. Hist. Soc. Northumb. 57: 
5-12. 

Latham, J. 1801. Supplement II to the General Synopsis of Birds. London. 

Latham, J. 1802. Supplementum indicts ornithologici sive systematis ornithologiae. London. 

Sharpe, R. B. 1890. Catalogue of the Passeriformes, or Perching Birds, in the Collection of 
the British Museum. Sturniformes. London. 

Whitehead, P. J. P. 1969. Zoological specimens from Captain Cook’s voyages. 7. Soc. 
Bibliog. Nat. Hist. 5: 161-201. 


Address: The Hancock Museum (Tyne and Wear Museums), Barras Bridge, Newcastle 
upon Tyne NE2 4PT, U.K. 


© British Ornithologists’ Club 1998 


C. B. Frith & D. W. Frith 7 Bull. B.O.C. 1998 118(1) 


Hybridization between Macgregor’s 
Bowerbird Amblyornis macgregoriae and the 


Streaked Bowerbird A. subalaris 
(Ptilonorhynchidae) of New Guinea 


by Clifford B. Frith S& Dawn W. Frith 
Received 4 December 1996 


The four species of ‘gardener’ bowerbirds of the genus Amblyornis 
(Ptilonorhynchidae) are confined to upland areas of New Guinea. All 
are known or assumed to be polygynous species in which males build 
elaborate ‘maypole’ type bowers and females alone construct and attend 
the nest and raise their offspring (Gilliard 1969, Diamond 1986). 

In all species individuals of both sexes are predominantly uniformly 
plumaged in buff, olive or rufous brown; and immature (crestless) 
males are like adult females. Adult males of three species are, however, 
adorned with a brilliant yellow (Golden-fronted Bowerbird A. 
flavifrons) to deep orange (Macgregor’s A. macgregoriae, Streaked 
Bowerbird A. subalaris) crest. That adult males of the fourth species 
(Vogelkop Bowerbird A. inornatus) lack a crest and are thus identical to 
their females yet build the most elaborate of all bowers led to the 
‘transferral effect’ theory of Gilliard (1969). This assumed that the 
forces of sexual selection (by females) have been transferred (or 
externalized) from morphological characters (the crest of adult males) 
to external objects (bowers) and that this ‘transferral’ has resulted in 
increasingly complex bower structures and their decoration. Thus as 
males of the various species have developed bower-building they have 
then simultaneously lost their crests to an extent correlated with the 
increasing complexity of their bower (Gilliard 1956, 1963, 1969). For 
full descriptions and illustrations of the birds and their bowers see 
Gilliard (1969), Cooper & Forshaw (1977) and Coates (1990). 

Crown colouration among the three monogamous non-bower- 
building bowerbirds or catbirds (Azluroedus spp.), and in the nine or 
ten members of the polygynous ‘avenue’-building genera Chlamydera 
and Sericulus, conspicuously reflects speciation in these groups 
(Gilliard 1969, Frith & Frith 1995, 1997a, 1997b). Similarly, both adult 
male crest colouration and, more particularly, crest length differ 
between taxa and reflect speciation within Amblyornis (Gilliard 1969, 
Diamond 1972, Schodde & McKean 1973, Pratt 1982, Frith & Frith 
1997b). 

Macgregor’s Bowerbird and the Streaked Bowerbird occur over 
predominantly different altitudinal ranges in eastern New Guinea. The 
higher range of 1600-3300 m) (occasionally down to 1200 m) asl of 
the former species is bordered for much of its southern and a third 
of its northern range by that of the latter species at 1000-1400 m 
(occasionally 700-1400 m) asl (Beehler et al. 1986)—see Figure 1. 


Collecting localities and altitudes of museum specimens of 


C. B. Frith & D. W. Frith 8 Bull. B.O.C. 1998 118(1) 


Figure 1. Map showing approximate distributions of Macgregor’s Bowerbird Amblyornis 
macgregoriae (vertical hatched areas) and Streaked Bowerbird A. subalaris (adjacent 
unhatched narrow strips) on the south-eastern Papua New Guinea peninsula. Inset map 
shows the New Guinea mainland with the vertical border line between Irian Jaya to its 
left and Papua New Guinea to its right with the area of the larger map solid black. 
Locations mentioned in the text are indicated by: AE=Mt. Albert Edward, AR=Aroa 
River, DD=Deva Deva, E=Efogi, I=Iola, M=Mafulu, MP=Murray Pass, 
SJR=Angabunga or St. Joseph River, T=Mt. Tafa, YI=Yule Island. Areas indicated by 
a question mark are those gardener bowerbirds remain to be recorded in. After Schodde 
& McKean (1973). 


Macgregor’s and Streaked Bowerbirds indicated the possibility of 
sympatry between the two in the northeastern part of the range of the 
latter species (Frith 1970). It was subsequently noted that collecting 
methods and data for the specimens concerned were not necessarily 
accurate (LeCroy 1971, Bell 1972). The latter author reported, 
however, that both species had been observed at Efogi at c. 1800 asl 
(Fig. 1), where Macgregor’s Bowerbird was in moss forest and the 
Streaked Bowerbird slightly below it. He wrote that “the species 
apparently need quite different habitats and their ranges apparently 
overlap merely because the moss-forest occurs at different altitudes in 
different areas with different climates’’. 

In a review of the taxonomy of the Amblyornis bowerbirds an 
Australian Museum specimen (AM O. 26264) was noted to possibly 
represent a hybrid individual between Macgregor’s and Streaked 
Bowerbirds (Schodde & McKean 1973). This specimen lacks a locality 
and has a 51mm long crest measured from its posterior base. 
Compared with Streaked Bowerbirds, it has a “‘rather heavy pale bill, 
rather pale-brown sides to the head, a rather dense crest, streaked 
throat and the following measurements (mm): wing 126; tail 82; tarsus 


C. B. Frith & D. W. Frith 9 Bull. B.O.C. 1998 118(1) 


36; bill (tip to nostril) 12.7. In these characters, it falls between A. 
subalaris and A. macgregoriae, and raises the possibility of occasional 
hybridization between the two species’ (Schodde & McKean 1973). 

This apparent hybrid was of interest as none had been authenticated 
within the bowerbird family notwithstanding a supposed unique 
intergeneric hybrid adult male specimen between the Satin Bowerbird 
Ptilonorhynchus violaceus and Regent Bowerbird Sericulus chryso- 
cephalus of eastern Australia (Marshall 1954, Gilliard 1969). While the 
authenticity of this supposed intergeneric hybrid combination is now 
seriously doubted, the unique specimen having long ago disappeared, 
an undoubted interspecific hybrid between the Great and Spotted 
Bowerbirds Chlamydera nuchalis x C. maculata of Australia has been 
described (Frith & Frith 1995). 


Methods 


While studying bird specimens at the American Museum of Natural 
History, New York (AMNH), we took the opportunity to examine 
skins of gardener bowerbirds for another study of this group (Frith & 
Frith 1997b). The collection contained an adult-plumaged specimen 
of Macgregor’s Bowerbird taken at Iola on 28 May 1929 and four 
adult-plumaged male Streaked Bowerbirds from the same locality 
during 1-5 June 1929, collected by Hannibal Hamlin of the Whitney 
South Sea Expedition. A Streaked Bowerbird specimen (AMNH 
831497) was recorded by Lee Crandall as taken at the adjacent village of 
Deva Deva at “‘4-5000’”’ (c. 1375 m) asl on 20 October 1928. Iola is a 
village, immediately east of Deva Deva and west and below Mafulu, on 
the mountain range just to the south of the Angabunga (or St Joseph) 
River, inland of Yule Island (Fig. 1). 

Iola and Deva Deva were also visited by the 1933-34 Archbold 
Expedition en route to Mt. Albert Edward via Mt. Tafa and the Murray 
Pass, but were stated to be only 700 m asl (Archbold & Rand 1935). 
These two gardener bowerbird species were also both collected by 
Hamlin and/or his collectors at Deva Deva. While acknowledging that 
Hamlin’s Macgregor’s Bowerbird locations are at approximately 700 m 
asl, Mayr & Rand (1937) noted they encountered the species only at 
2000-2800 m asl. As 770m is well below even the extreme lowest 
altitude normally frequented by Macgregor’s Bowerbird it seems likely 
Hamlin and his collectors climbed to procure the two specimens of that 
species or obtained them from Papuans that did so for him. 

We examined and measured all gardener bowerbird specimens from 
Iola and immediately adjacent localities (Table 1). All wing and tail 
measurements were made with the same rulers, all others with the same 
electrical digital callipers, and all by CBF. Wing length measured was 
the flattened and straightened, thus maximised, chord using a stopped 
steel rule. Tail length was measured from the point of entry of the 
central pair into the bird’s skin to the tip of the longest feather with a 
small steel rule. Bill length was measured from the bill tip to the 
cranio-facial hinge. Tarsus length was that of the tarsometatarsal bone. 
One or two crest length measurements were taken of adult males, the 


Bull. B.O.C. 1998 118(1) 


10 


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length from posterior base (cf. Schodde & McKean 1973) and that of 
the exposed or visible length of the orange crest (‘Table 1). 


Results and discussion 


Of the four AMNH Iola specimens identified as Streaked Bowerbirds, 
three had fully developed orange crest plumes marked with small 
amounts of deep brown, from fine short flecking (AMNH 330488) to 
numerous larger streakings (330491). While the identity of these four 
specimens has never been questioned, it was noted by Mayr & Rand 
(1937) that one of them had the usually pronounced throat and breast 
streaking much reduced. The other three AMNH Iola males have a 
crest bordered at the outer edges with a contrastingly dark, almost a 
reddish-black, brown—a feature characteristic of Streaked Bowerbirds, 
present in the three adult males from nearby Deva Deva. Adult male 
Macgregor’s do have brown feathering at the outer border of their crest 
but it is far less dark and discrete than in Streaked Bowerbirds, being 
much the same olive-brown as the rest of their head and nape. 

The fourth adult male supposed Streaked Bowerbird (AMNH 
330487) from Iola does not exhibit the dark crest border typical of the 
species but has a crest intermediate in colour and tone between that 
species and the Iola Macgregor’s Bowerbird. The sides of its face 
are not lighter than in Streaked Bowerbirds from Iola. It differs 
conspicuously from adult male Streaked Bowerbirds from there and 
elsewhere in having only the slightest trace of throat and chest 
streaking, notwithstanding known variation of ventral markings in the 
Streaked Bowerbird (cf. Schodde & McKean 1973). We view this 
‘masking’ of typical Streaked ventral streaking as an expression of the 
influence of ventrally plain Macgregor’s Bowerbird genes. The base of 
the lower mandible of the apparent hybrid individual is paler than that 
of most Streaked Bowerbird specimens, only that of specimen AMNH 
330490 being similarly pale. The hybrid specimen (AMNH 330487) 
unfortunately lacks its tail. 

All three Iola adult male Streaked Bowerbirds of normal appearance 
have a crest shorter than 40 mm in length, measured from the posterior 
base. Crest length in 12 AMNH adult male Owen Stanley Mountains 
Streaked Bowerbirds, additional to those detailed in Table 1, average 
35 (range 24—-40)mm. That of eight other adult male Streaked 
Bowerbirds, in Australian collections, average 39 (range 37-42) mm 
(Schodde & McKean 1973). The latter authors also cite crest lengths 
from posterior base of three adult male Macgregor’s Bowerbirds from 
the western Owen Stanley Range of 52, 57 and 59mm. Thus the 
72mm exposed crest length of the Iola hybrid specimen AMNH 
330487 is intermediate between the mutually exclusive crest lengths of 
Streaked and Macgregor’s Bowerbirds. 

The hybrid specimen was recorded as collected at Iolaon 1 June, the 
three Streaked Bowerbirds on 4-5 June and the Macgregor’s 
Bowerbird on 28 May. We therefore see the former’s larger wing 
length, conspicuously longer crest and lack of ventral streaking, 
compared with these features of Streaked Bowerbirds from the same 


C. B. Fnth & D.W. Frith 12 Bull. B.O.C. 1998 118(1) 


Figure 2. Dorsal appearance of AMNH specimens of adult males of: Amblyornis 
macgregoriae (33048), A hybrid A. macgregoriae X A. subalaris (330487, lacking tail) and 
three A. subalaris (330488, 330490, 330491) seen from top to bottom respectively. Note 
intermediate crest length and dorsal markings of the hybrid. 


area (‘Table 1), as too great a coincidence to be indicative of intraspecific 
variation. Hence we consider it an A. macgregoriae X A. subalaris 
hybrid individual exhibiting stronger evidence of the genes of both 
putative parents than the specimen (AM O. 26264) of unknown locality 
thought also to represent this hybrid combination (Schodde & 
McKean 1973). Our measurements of both crest from posterior base 
and exposed crest support this conclusion and are consistent with 
measurements of both putative parent species presented by Diamond 


C. B. Frith & D. W. Frith 13 Bull. B.O.C. 1998 118(1) 


(1972) and Schodde & McKean (1973). We suppose it was this AMNH 
hybrid specimen that Mayr & Rand (1937) noted as being less typical of 
Streaked Bowerbirds (see above). 

Male gardener bowerbirds probably require at least five or six years — 
to acquire adult plumage, as other sexually dimorphic bowerbird 
species are known to do (Disney 1970, Disney & Lane 1971, Vellenga 
1980, Frith & Frith unpubl. data). Adult-plumaged male gardener 
bowerbirds would therefore represent but a small proportion of their 
species populations. It can therefore be appreciated that hybrid 
individuals in female (and immature male) plumage would be difficult 
to identify from their appearance in museum specimens. Genetic 
‘fingerprinting’ studies in areas where the two species may meet clearly 
have the potential to prove rewarding as it would seem that 
hybridization may be more common in the family than has been 
previously conceded (Frith & Frith 1995). 

Macgregor’s and Streaked Bowerbirds are for the most part 
separated by mutually exclusive altitudinal ranges and associated 
differences in habitat. While specimens demonstrate that the two 
species occasionally hybridize in the wild, providing clear evidence that 
they are at some places sympatric (sensu stricto cf. Amadon & Short 
1992), they are better described as parapatric (Mayr 1963: 672, Wilson 
1975, Kemp 1995). Specimens in the AMNH show this is clearly the 
case in the Iola and Deva Deva area, and it may well be the case 
elsewhere, potentially including the upper Aroa River (Frith 1970). It 
remains to be assessed whether parapatry occurs within natural 
altitudinal ranges/habitats, or if it may only result from habitat 
modification caused by agriculture (Gilliard in Mayr & Gilliard 1952, 
Gilliard 1959) or more severe (geological) habitat destruction. 


Acknowledgements 


We are grateful to the authorities of the American Museum of Natural History for access 
to specimens, to the Chapman Fund of that institution for support and to Mary LeCroy 
for kind hospitality and much help. The Field Museum of Natural History, Chicago sent 
a specimen loan to New York for us and we thank David Willard for arranging this. Mary 
LeCroy and Richard Schodde provided thoughtful and constructive comment on a draft 
of this contribution, which we dedicate to the memory of Harry L. Bell. 


References: 
Amadon, D. & Short, L. L. 1992. Taxonomy of lower catergories—suggested guidelines. 
Bull. Br. Orn. Cl., Centenary Suppl. 112A: 11-38. 
Archbold, R. & Rand, A. L. 1935. Results of the Archbold Expeditions No. 7. Summary 
of the 1933-1934 Papuan Expedition. Bull. Amer. Mus. Nat. Hist. 68: 527-579. 
Beehler, B. M., Pratt, T. K. & Zimmerman, D. A. 1986. Birds of New Guinea. Princeton 
Univ. Press. 

Bell, H. L. 1972. Altitudinal distribution of bowerbirds of the genus Amblyornis. Emu 72: 
34. 

Coates. B. J. 1990. The Birds of Papua New Guinea Vol. 2. Dove, Alderley. 

gh W. T. & Forshaw, J. M. 1977. The Birds of Paradise and Bower Birds. Collins, 

ydney. 

Diamond, J. M. 1972. Avifauna of the Eastern Highlands of New Guinea. Publ. Nuttall 
Orn. Club 12. 

Diamond, J. M. 1986. Biology of birds of paradise and bowerbirds. Ann. Rev. Ecol. Syst. 
17: 17-37. 


G. N. Mauricio & R. A. Dias 14 Bull. B.O.C. 1998 118(1) 


Disney, H. J. de S. 1971. Bird in the hand—Regent Bowerbird Sericulus chrysocephalus. 
Austral. Bird Bander 9: 14-15. 

Disney, H. J. de S. & Lane, S. G 1971. Moult, plumage and banding of the Regent 
Bowerbird. Austral. Bird Bander 9: 11-13. 

Frith, C. B. 1970. Sympatry of Amblyornis subalaris and A. macgregoriae in New Guinea. 
Emu 70: 196-197. 

Frith, C. B. & Frith, D. W. 1995. Hybridization between the Great and Spotted 
Bowerbird Chlamydera nuchalis and C. maculata: an authenticated hybrid bowerbird 
(Ptilonorhynchidae). Mem. Queensland Mus. 38: 471-476. 

Frith, C. B. & Frith, D. W. 1997a. Chlamydera guttata carteri Mathews 1920—an 
overlooked subspecies of Western Bowerbird (Ptilonorhynchidae) from North West 
Cape, Western Australia. Rec. West. Aust. Mus. 18: 219-226. 

Frith, C. B. & Frith, D. W. 1997b. A distinctive new subspecies of Macgregor’s 
Bowerbird (Ptilonorhynchidae) of New Guinea. Bull. Brit. Orn. Cl. 117: 199-205. 

Gilliard, E. T. 1956.. Bower ornamentation versus plumage characters in bower-birds. 
Auk.73:.450-451. 

Gilliard, E. T. 1959. The ecology of hybridization in New Guinea Honeyeaters (Aves). 
Amer. Mus. Novitates 1937 1-26. 

Gilliard, E. T. 1963. The evolution of Bowerbirds. Scientific American 209: 38-46. 

Gilliard, E. T. 1969. Birds of Paradise and Bower Birds. Weidenfeld & Nicholson, 
London. 

Kemp, A. 1995. The Hornbills. Oxford Uni. Press, Oxford. 

LeCroy, M. K. 1971. Sympatry in bowerbirds of genus Amblyornis. Emu 71: 143. 

Marshall, A. J. 1954. Bower-Birds: their displays and breeding cycles Oxford Uni. Press, 
Oxford. 

Mayr, E. 1963. Animal Species and Evolution. Belknap Press, Harvard Uni. Press, 
Cambridge, Mass. 

Mayr, E. & Gilliard, E. T. 1952. The Ribbon-tailed Bird of Paradise (Astrapia mayeri) 
and its allies. Amer. Mus. Novitates 1551: 1-13. 

Mayr, E. & Rand, A. L. 1937. Results of the Archbold Expeditions. 14. Birds of the 
1933-1934 Papuan Expedition. Bull. Amer. Mus. Nat. Hist. 73: 1 — 248. 

Pratt, T. K. 1982. Additions to the avifauna of the Adelbert Range, Papua New Guinea. 
Emu 82: 117-125. 

Schodde, R. & McKean, J. L. 1973. Distribution, taxonomy and evolution of the 
gardener bowerbirds Amblyornis spp. in eastern New Guinea with descriptions of 
two new subspecies. Emu 73: 51-60. 

Vallenga, R. E. 1980. Moults of the Satin Bowerbird Ptilonorhynchus violaceus. Emu 80: 
49-54. 

Wilson, E. O. 1975. Sociobiology: The new synthesis. Belknap Press, Harvard Uni. Press, 
Cambridge, Mass. 


Address: ‘Prionodura’, P.O. Box 581, Malanda, Queensland 4885, Australia. 


© British Ornithologists’ Club 1997 


Range extensions and new records for forest 
birds in southern Rio Grande do Sul, Brazil 


by Giovanni N. Mauricio & Rafael A. Dias 
Received 23 December 1996 
The southernmost distributional limits of forest birds in southern 


Brazil are particularly evident in the State of Rio Grande do Sul, where 
the Atlantic Forest has its last domains. Distributional patterns of 


G. N. Mauricio & R. A. Dias 15 Bull. B.O.C. 1998 118(1) 


RIO URUGUAI 


RIO PIRATINI a 


Figure 1. Map showing localities mentioned in text. 1, Turvo state reserve; 2, Nonoai; 3, 
Erebango; 4, Sananduva; 5, Lagoa Vermelha; 6, Passo Fundo; 7, Panambi (Nova 
Wurtemburg); 8, Santo Angelo; 9, Garruchos; 10, Itaqui; 11, Bom Jesus; 12, Aparados da 
Serra national park; 13, Farroupilha; 14, Torres; 15, Sado Francisco de Paula; 16, Barra do 
Ouro; 17, Osorio; 18, Arroio Grande; 19, Taquara; 20, Campo Bom; 21, Sapucaia do Sul; 
22, Novo Hamburgo (Hamburgo Velho); 23, Sapiranga; 24, Sao Sebastido do Cai; 25, 
Montenegro (Monte Negro); 26, Poco das Antas; 27, Cruzeiro do Sul; 28, Santa Cruz do 
Sul; 29, Santa Maria; 30, Triunfo; 31, Guaiba (Pedras Brancas); 32, Porto Alegre; 33, 
Viamao; 34, Encruzilhada do Sul; 35, Rincao dos Pereira; 36, Santana da Boa Vista; 37, 
Camaqua; 38, Sao Lourenco (do Sul); 39, Cangucu; 40, Pinheiro Machado; 41, Pelotas; 
42, Herval; 43, Arroio Grande; 44, Taquaral; 45, Arroio do Padre; 46, Arroio Andrade; 
47, Parque Farroupilha; 48, Rincao da Caneleira; 49, Arroio Cadeia; 50, Arroio dos 
Porcos; 51, Santa Eulalia; 52, Santo Amor; 53, Arroio Moreira; 54, Arroio Padre Doutor; 
55, Cerro das Almas; 56, UFPEL botanical garden; 57, Pontal da Barra (on Laguna dos 
Patos littoral); 58, Rio Piratini. I, Serra do Herval; II, Serra dos Tapes. The dotted line 
represents the Serra Geral escarpment. 


forest birds in Rio Grande do Sul identified by Belton (1984, 1985) 
indicate distinct forested areas as limits for some groups of species. In 
this way, the northeastern littoral coastal forests are considered the 
southern limit for some lowland species (e.g. Formicarius colma, 


G. N. Mauricio & R. A. Dias 16 Bull. B.O.C. 1998 118(1) 


Myrmotherula unicolor, Platyrinchus leucoryphus), whereas the northern 
forests, along the Rio Uruguai, constitute limits for other ones (e.g. 
Pteroglossus castanotis, Drymophila rubricollis, Colonia colonus) which 
apparently reach the state from inland Brazil. The forests of the 
escarpment of the Serra Geral highlands, marking the transition 
between the forested northern half of Rio Grande do Sul and the 
more open south, although seemingly being the southern limit for 
predominantly montane species (e.g. Chamaeza ruficauda, Hylopezus 
nattererl, Piprites pileatus) do not show a precise effect on bird 
distribution, as Belton (1984: 391) states: ““‘Most species with ranges to 
the south and the west are limited by this barrier, but many from the 
north are not affected by it, either not reaching it in significant numbers 
or, if they do, passing beyond it into the central trough or southeastern 
hills’’. The presence of forest birds south of the escarpment is mostly 
related to the occurrence of a formerly continuous forest block on the 
eastern slopes of Rio Grande do Sul’s southeastern hills (Serra dos 
‘Tapes and Serra do Herval). This forest unit, poorly studied and 
virtually unknown to ornithology, is a semideciduous forest of humid 
climate (Floresta Estacional Semi-Decidual; Brasil 1986). Therefore, 
although the southern half of Rio Grande do Sul is typically grassland, 
an appendix of the Atlantic Forest (sensu lato) extends south of the 
escarpment and the adjacent central trough (i.e. south of 30°S to 
approximately 31°40’'S), covering the humid slopes of the eastern 
watershed of Rio Grande do Sul’s Crystalline Shield (southeastern 
hills), a deeply eroded (medium height 300m) prolongation of 
southeastern Brazil’s Serra do Mar mountains. 

Ihering (1899) was the first author to mention the few forest birds 
known from this region, citing specimens obtained in the municipalities 
of Sao Lourencgo (31°22’S, 51°58’W, now Sao Lourenco do Sul) in the 
1880s and Pedras Brancas, the former name of Guaiba (30°07’S, 
51°20’W), a town on the northeastern limit of this forest. This author 
also contested the origin Pelotas (31°45’S, 52°20’W; a city on this 
forest’s southern limit) of specimens from the British Museum’s Joyner 
collection, with which Belton (1984, 1985) agreed. Pinto (1938) 
reported specimens collected in Sao Lourenco around 1900, probably 
obtained within the limits of the semideciduous forest. E. Kaempfer 
did not collect in Sao Lourenco, only along the Laguna dos Patos 
littoral, as stated by Naumburg (1935), but also in the higher part of the 
municipality (see text on the Hooded Berry-eater Carpornis cucullatus). 
More recently Belton (1984, 1985, 1994) mentions additional forest 
birds from the southeastern hills, as well as records of other birds from 
localities already cited above, without however relating their presence 
to the occurrence of a previously forested area, nowadays extremely 
fragmented. 

Between October 1993 and July 1996, we conducted an ornithologi- 
cal survey in several forest remnants located in the southernmost 
domains of this former forest region, documenting with tape recordings 
new records of forest birds that represent substantial southward range 
extensions (some of these are sight records). In the municipality of 
Pelotas, 6 forest remnants were studied: Arroio Andrade (31°27'S, 


G. N. Mauricio & R. A. Dias 17 Bull. B.O.C. 1998 118(1) 


52°28’W; alt. 300m, 150ha of primary forest); Arroio do Padre 
(31°24’S, 52°23'W; alt. 160-180 m, 10 ha of primary forest); Rincao da 
Caneleira (31°31'S, 52°35’W; alt. 200m, 10ha of primary forest); 
Parque Farroupilha (31°30'S, 52°34'W; alt. 160m, 10 ha of primary 
forest); Arroio dos Porcos (31°33’S, 52°32'W; alt. 60m, 15 ha of 
primary forest) and Santa Eulalia (31°34’S, 52°32'W; alt. 100 m, 20 ha 
of primary forest). On the border of the municipalities of Pelotas and 
Morro Redondo we visited Arroio Cadeia (31°33’S, 52°34'W; alt. 
100-120 m, 50ha of primary forest) and Santo Amor (31°40’S, 
52°35'W; alt. 100 m, small fragments of mixed primary and secondary 
growths). In the municipality of SAo Lourenco the only remnant visited 
was Taquaral (31°20'S, 52°23’W; alt. 100-200 m, 200 ha of primary 
forest). All these forest remnants were sporadically visited during the 
years, with the exception of Arroio Andrade, visited 8 times (covering 
22 days) between February 1995 and May 1996. Additional work was 
done outside the borders of the original forest (sensu Brasil 1986), on 
gallery forests and on an isolated 500 ha forest block at Cerro das Almas 
(31°46’S, 52°35’W), a small chain of granitic hills with altitudes varying 
from 100 to 250 m located in the municipality of Capado do Ledo. 


SHORT-TAILED HAWK Buteo brachyurus 


Ihering (1899) and Gliesch (1930) recorded this hawk in Rio Grande 
do Sul at Taquara and Porto Alegre, respectively. More recently, E. 
Willis observed this species in three different locations along the 
southern escarpment, while W. Voss and L. R. C. da Silva recorded it 
twice at Sapucaia do Sul and W. Voss saw a pair in apparent courtship 
behaviour at Barra do Ouro (Belton 1984). Additionally, W. Voss and 
others observed this species at Triunfo and Sao Sebastiao do Cai, and 
G. Bencke found a dead individual at Cruzeiro do Sul (Belton 1994). 
We found B. brachyurus as resident at Arroio Andrade, with up to three 
individuals recorded in a day, including immatures (observed on 17 
February 1996) and black morphs. An immature was seen at Rincdao da 
Caneleira in late January and early February 1995. Two immatures and 
a black adult were observed at Arroio Cadeia on several days in 
February 1995. Also recorded at Cerro das Almas and an individual 
was seen flying over nearby Arroio Moreira gallery forest (31°43’S, 
52°28’W) on 16 June 1996. These records represent a southward 
distributional extension of c. 200 km from the southern range limit (1.e. 
30°S) marked by Belton (1994). 


BARRED FOREST-FALCON Micrastur ruficollis 


Ihering (1899), Gliesch (1930), Pinto (1938) and Camargo (1962) 
mentioned specimens from 'Taquara, Porto Alegre, Sao Lourengo and 
Santa Maria, respectively. Besides these, Belton (1984) cites records 
from scattered localities in the northern highlands and one from the 
southeastern hills. We found this species throughout the year at Arroio 
Andrade and also at Santa Eulalia on 23 November 1996 and at 
Taquaral on 5 November 1995. The nearest record (from Sao 
Lourengo, just north of the study area) is given by Pinto (1938), based 
on a specimen collected in 1900. 


G. N. Mauricio & R. A. Dias 18 Bull. B.O.C. 1998 118(1) 


RUDDY QUAIL-DOVE Geotrygon montana 

Known in Rio Grande do Sul from scattered points in the state’s 
northern and northeastern sectors and from the central portion of the 
southeastern hills, north of the Rio Camaqua (Belton 1984). Mentioned 
from Taquara by Ihering (1899) and Farroupilha and Osorio by 
Camargo (1962). On 1 February 1995 we observed a female sitting on a 
nest with two chicks at Rincao da Caneleira, and on 21 February 1995 
we saw a male at Arroio Andrade. These records extend the known 
distribution of this species c. 100 km to the south. 


RUSTY-BARRED OWL Strix hylophila 

We observed and tape-recorded up to three individuals of this owl on 
numerous occasions throughout the year in three small forest remnants 
(Parque Farroupilha, Rincao da Canaleira and Arroio dos Porcos). It is 
more numerous at Arroio Andrade, with up to six individuals detected. 
Found by Belton (1984) in northeastern Rio Grande do Sul, primarily 
above the escarpment and once in the southeastern hills, near Santana 
da Boa Vista. Ihering (1899) cited this species from Taquara and Pinto 
(1938) mentioned a specimen from Sao Lourenco. 


SEMI-COLLARED NIGHTHAWK Lurocalis semitorquatus 

Mentioned for T’aquara by Ihering (1899) and Poco das Antas by 
Gliesch (1930). Belton (1984) reported it as common near the 
escarpment and in the extreme north, along the Rio Uruguai, with 
an isolated record for the central highlands and another for the 
southeastern hills, north of the Rio Camaqua. This species was 
recorded in November, December and February at Rincado da 
Canaleira, Arroio Cadeia and Arroio Andrade, which extends the 
known distribution of this species 100 km southwards. 


ASHY-TAILED SWIFT Chaetura andrei 

First mentioned for Rio Grande do Sul by Camargo (1962), based on 
a specimen collected in Farroupilha. Belton (1984) recorded this species 
from the state’s northeastern quarter (except the highest areas) and 
along the escarpment. At Arroio Andrade we observed this swift 
regularly between October and late March, disappearing from the area 
in the fall and winter. Some individuals were additionally seen at 
‘Taquaral on 5 November 1996. These records represent a southward 
extension of the known distribution of this species by c. 200 km. 


RED-BREASTED TOUCAN Ramphastos dicolorus 


Reported from Taquara by Ihering (1899) and Poco das Antas by 
Gliesch (1930). Pinto (1938) and Camargo (1962) mentioned specimens 
collected respectively at Itaqui (on the Rio Uruguai) and Passo Fundo. 
According to Belton (1984), the distribution of R. dicolorus in Rio 
Grande do Sul follows the escarpment, from Santa Maria to Aparados 
da Serra national park and also in the extreme and central north, 
disregarding the specimen exhibited at Camaqua’s Museu de Aves 
Empalhadas, collected at Encruzilhada do Sul, a town in the northern 
sector of the southeastern hills. Later, Belton (1994) admits the bird’s 


G. N. Mauricio S R. A. Dias 19 Bull. B.O.C. 1998 118(1) 


casual occurrence in the southeastern hills, based on G. Bencke’s 
communication of an individual killed at Encruzilhada do Sul. Haffer 
(1974), in addition to the localities mentioned above in the northern 
half of Rio Grande do Sul, marks an additional point south of 32°S 
(in the municipality of Arroio Grande; 32°15’'S, 53°05’W) in his 
distribution map of R. dicolorus. Haffer (in litt. 1996) informs us that 
this point refers to Berlepsch & Ihering’s (1885) citation of Arroio 
Grande (29°45’S, 50°45’W), a locality near Taquara, and not to the 
southern village of the same name mistakenly shown on the map. In our 
study area, this species is resident and particularly numerous at Arroio 
Andrade (groups of up to 20 individuals), where a pair was also seen 
feeding two young in a cavity 8 m up in an Alchornea triplinervea tree, 
on 22 February 1995. Pairs were also observed at Rincao da Caneleira 
and Parque Farroupilha. Additionally, there is a specimen at 
Universidade Catolica de Pelotas found dead on a highway near Rincao 
da Caneleira on 14 June 1994. 


WHITE-BROWED WOODPECKER Piculus aurulentus 

Cited for Taquara by Ihering (1899) and Poco das Antas and Porto 
Alegre by Gliesch (1930). Specimens are mentioned from Itaqui by 
Pinto (1938) and Bom Jesus and Passo Fundo by Camargo (1962). 
Belton (1984) found this woodpecker in the state’s northeastern quarter 
and mentioned two isolated records from the extreme north and the 
southeastern hills (one north and another south of the Rio Camaqua). 
We recorded this species in Santo Amor, Rincado da Caneleira and 
Arroio Andrade, on 25 May 1994, 19 March 1995 and 10 October 1995, 
respectively. 


LINEATED WOODPECKER Dryocopus lineatus 


Ihering (1899) mentioned this species from 'Taquara and Gliesch 
(1930) from Poco das Antas and Santo Angelo. Subsequently recorded 
from Bom Jesus (Camargo 1962). Belton (1984) found it in the north, 
northeast and northwestern sectors of the state and reported it from the 
southeastern hills, with records from Rincao dos Pereira and Sao 
Lourengo, the latter based on a specimen cited by Pinto (1938). A pair 
were attending a nest with two young, 3 m up in a dead tree at Parque 
Farroupilha on 2 November 1993 and later, in the same locality, a 
female was recorded on 25 June 1994. An individual was observed at 
Arroio Andrade on 27 March 1995. 


ROBUST WOODPECKER Campephilus robustus 


We found this species only at Arroio Andrade, where a small family 
party (adult pair and young female) was initially observed on 21 
February 1995. Isolated individuals were later seen during the year and 
the female’s voice was recorded on two occasions. The nearest previous 
records are for the foot of the escarpment, at Taquara (Ihering 1899) 
and Poco das Antas (Gliesch 1930), c. 250 km to the north. Additionally 
mentioned for Santo Angelo by Gliesch (1930) and Bom Jesus by 
Camargo (1962). Belton (1984) cited scattered records from the high 
northeast, extreme north and Garruchos, considering this species rare. 


G. N. Mauricio & R. A. Dias 20 Bull. B.O.C. 1998 118(1) 


WHITE-THROATED WOODCREEPER X7iphocolaptes albicollis 

Common resident at Arroio Andrade, the only forest remnant where 
it was found. Belton (1994) marked its distribution north of 30°S, 
doubting the origin Camaqua of the specimen at Museu de Aves 
Empalhadas of Camaqua and, despite G. Bencke’s voice-based winter 
record from Santa Cruz do Sul, in August 1990, considers that this 
species is possibly a summer resident in Rio Grande do Sul, which is 
not confirmed by our records. Previously mentioned from sparse 
localities in the northern half of Rio Grande do Sul: Taquara (Ihering 
1899), Itaqui (Pinto 1938), Bom Jesus and Osorio (Camargo 1962). 


PLANALTO WOODCREEPER Dendrocolaptes platyrostris 

We found this species in 7 forest remnants (Rincao. da Caneleira, 
Parque Farroupilha, Arroio Andrade, Arroio dos Porcos, Arroio Cadeia 
and Santa Eulalia), both in primary and in old and recent secondary 
growths. Also recorded further south, outside the limits of the 
originally forested region, in gallery forests along the Rio Piratini 
(31°51’S, 52°50’W). According to Belton (1984), the distribution of this 
species in Rio Grande do Sul covers primarily the northern half of the 
state, with only one record from the southeastern hills, just south of 
the Rio Camaqua. Mentioned from 'Taquara (Ihering 1899), Itaqui 
(Pinto 1938), Santo Angelo and Porto Alegre (Gliesch 1930). 


RUFOUS-CAPPED SPINETAIL Synallaxis ruficapilla 

Known in Rio Grande do Sul north from 30°S as far as Santa Maria 
to the west and west across the state north of 28°30’S (Belton 1984). 
Besides citing it from T’aquara, Ihering (1899) mentioned a record 
south of 30°S, from Pedras Brancas, overlooked by modern authors. 
Pinto (1938) recorded this spinetail from Nova Wurtemburg (now 
Panambi) and Camargo (1962) from Farroupilha. Recorded regularly at 
Arroio Andrade, primarily along a clearing with dense undergrowth, 
where we tape-recorded it twice, and also in a bamboo thicket. 
Additionally found at Cerro das Almas. Our records represent a 
southward range extension of c. 150 km from the historical record of 
Pedras Brancas. 


SHORT-TAILED ANTTHRUSH Chamaeza campanisona 

Recorded by Belton (1985) in forests along the escarpment, central 
trough (east of Santa Maria) and in the northern sector above 28°30'S. 
Previously known from Taquara, Pedras Brancas (Ihering 1899), Porto 
Alegre (Gliesch 1930), Novo Hamburgo (Pinto 1938), Erebango, 
Sananduva, Sapiranga (Naumburg 1939) and Santa Maria (Camargo 
1962). A recent record from gallery forests of the Rio Camaqua in the 
municipality of Cangucu in the southeastern hills is given by G. Bencke 
(Belton 1994). This species was recorded only in the larger forest 
fragments (Arroio Andrade and ‘Taquaral), being usually heard 
throughout the year. 


RUFOUS GNATEATER Conopophaga lineata 


This species was recorded in 6 forest remnants of our study area in 
the southeastern hills (Rincao da Caneleira, Parque Farroupilha, Arroio 


G. N. Mauricio S R. A. Dias 21 Bull. B.O.C. 1998 118(1) 


Andrade, Arroio dos Porcos, Arroio Cadeia and Santo Amor), being an 
uncommon resident. Also found at sea level in swampy forests of Pontal 
da Barra marsh (31°47'S, 52°14’W) and nearby sites along Laguna dos 
Patos in the municipality of Pelotas, where this gnateater is a scarce 
resident. Previously recorded in Rio Grande do Sul from localities in 
the state’s northern half. Ihering (1899) mentioned it from T’aquara 
and Gliesch (1930) from Porto Alegre. Naumburg (1937) reported 
specimens collected by E. Kaempfer at Hamburgo Velho (nowadays 
Novo Hamburgo), Sananduva, Nonoai, Sapiranga and Santa Cruz. 
Pinto (1938) cited it from Novo Hamburgo and Camargo (1962) from 
Farroupilha, Porto Alegre and Viamao. According to Belton (1985), the 
distribution of C. lineata in the state is restricted to the area north of 
the central trough, east of 54°W, with sparse records to the west. Our 
records represent a southward extension of the known distribution of 
this species by c. 200 km. 


MOUSE-COLOURED TAPACULO Scytalopus speluncae 


This species was recorded in all studied forest remnants; it is 
particularly abundant at Cerro das Almas, with up to twelve individuals 
recorded in a forest patch of less than 5 ha. Occurs at altitudes of about 
10 m above sea level near Cerro das Almas and in Arroio Padre Doutor 
gallery forest (31°44’S, 52°29’W). First mentioned from Rio Grande do 
Sul by Camargo (1962), based on specimens collected in Bom Jesus 
and Farroupilha. Belton (1985) recorded S. speluncae only in the state’s 
extreme north and along the eastern portion of the top of the 
escarpment, and also found an individual near Pinheiro Machado, c. 
100 km westward of our study site. 


PLANALTO TYRANNULET Phyllomyias fasciatus 


This summer resident is mentioned from the vicinity of the 
northeastern escarpment and the central part of the state, from the 
Santa Catarina border to the central trough, north of 30°S, also with a 
record from the southeastern hills near Santana da Boa Vista, where it 
was found by E. Willis (Belton 1985). Previously known in Rio Grande 
do Sul only from Taquara (Ihering 1899). We found this species most 
frequently at Arroio Andrade and also at Rincado da Caneleira, Arroio 
do Padre and Taquaral remnants, always in spring and summer. These 


records represent a southward extension of c. 100 km from the site of 
E. Willis’s record. 


ROUGH-LEGGED TYRANNULET Phyllomyias burmeisteri 


An unseen bird calling in the canopy of Arroio Cadeia forest was 
tape-recorded on 3 September 1995 and later identified by José 
Fernando Pacheco and Bret Whitney as this species. Later heard and 
tape-recorded at Arroio Andrade on 9 October 1995 and 12 May 1996. 
Belton (1985) recorded this species seven times in scattered localities of 
Rio Grande do Sul, north of 29°S (extreme north, northeast and 
northwest). Our records represent a southward extension of c. 300 km 
in the known distribution of P. burmeistert. 


G. N. Mauricio S R.A. Dias 22 Bull. B.O.C. 1998 118(1) 


GREENISH TYRANNULET Phyllomyias virescens 

We usually found only one pair of this species in the smaller forest 
remnants studied (Rincado da Caneleira, Arroio Cadeia, Arroio dos 
Porcos and Santo Amor) and up to four pairs in the larger ones (Arroio 
Andrade and Taquaral), regularly tape-recording its voice. Belton 
(1985) cited only four records from Rio Grande do Sul, three from the 
top of the escarpment in the northeast and one from Garruchos, in the 
extreme west, and stated that Kaempfer collected 13 specimens from 
the north-central sector in 1928-1929. The nearest previous record is 
located c. 300 km to the north of our study area. 


YELLOW-OLIVE FLYCATCHER To/momyias sulphurescens 


We regularly recorded this flycatcher in all forest fragments studied, 
including Cerro das Almas. Also found in gallery forests of Rio Piratini 
and Arroio Moreira and occasionally in forest patches on the coastal 
plain, at the botanical garden of Universidade Federal de Pelotas 
(31°48'S, 52°25'W). Previously known in Rio Grande do Sul north of 
the central trough (Belton 1985), including Porto Alegre (Gliesch 1930, 
Camargo 1962), Sao Francisco de Paula, Campo Bom, Sananduva, 
Sapiranga, Lagoa Vermelha and Nova Wurtemburg (Pinto 1944). 


WHITE-THROATED SPADEBILL Platyrinchus mystaceus 

Mentioned for Taquara (Ihering 1899), Farroupilha (Camargo 1962), 
Novo Hamburgo and Monte Negro (Pinto 1944). Besides these 
localities, Belton (1985) included in his proposed distribution of 
P. mystaceus in Rio Grande do Sul the state’s extreme north and 
northeastern sectors, with one record for Garruchos. Common in the 
larger forest fragments studied (Arroio Andrade and Taquaral), less so 
in the smaller ones (Rincado da Caneleira, Arroio Cadeia, Santa Eulalia 
and Santo Amor). These records extend the known distribution of this 
species by c. 200 km southwards. 


GREENISH SCHIFFORNIS Schiffornis virescens 

According to Belton (1985), in Rio Grande do Sul this species is a 
common resident in forests of the extreme north close to the Rio 
Pelotas and Rio Uruguai downstream to Garruchos, and uncommon in 
the vicinity of the escarpment from Torres to 52°30’W. Mentioned for 
Taquara by Ihering (1899). We found and tape-recorded isolated 
individuals of this species on two occasions (22 March and 10 October 
1995) at Arroio Andrade. More common at Taquaral forest, where we 
recorded more than six territorial individuals in 5 November 1995. Our 
records extend this species’ known distribution by c. 200 km to the 
south. 


HOODED BERRYEATER Carpornis cucullatus 

We found this endemic cotingid of the Brazilian Atlantic Forest in 
six forest remnants southwards to 31°34’S. In the smaller fragments 
(Rincao da Caneleira, Parque Farroupilha, Arroio do Padre and Santa 
Eulalia) only two individuals were recorded, whereas at Arroio Cadeia 
four and at Arroio Andrade up to eight singing males were detected. 


G. N. Mauricio GS R. A. Dias 23 Bull. B.O.C. 1998 118(1) 


Previously mentioned in Rio Grande do Sul from Taquara and Pedras 
Brancas by Ihering (1899), Osorio and Viamao by Camargo (1962) and 
Poco das Antas and Porto Alegre by Gliesch (1930). Belton (1985) 
limited the distribution of C. cucullatus between the northern littoral 
and the eastern end of the central trough (an area that covers the above 
mentioned localities) and additionally cited that Kaempfer found it in 
the southeastern hills, marking on his species map (Belton 1985: 89) 
two points south of 31°S without precise localities. Belton (zn litt. 1995) 
informs us that one of the points (north of Pelotas) represents a site 
west of Sao Lourenco, at 122 m altitude, not mentioned by Naumburg 
(1935), where two specimens (nos. 321984 and 321985 housed in the 
American Museum of Natural History) were collected by Kaempfer on 
14-15 October 1931, and that the other one (southwest of Pelotas) 
represents I[hering’s (1899) citation for Serra do Herval. However, the 
Serra do Herval where Ihering heard this species is located north of 
31°S, being the eastern watershed of the southeastern hills between the 
Rio Camaqua and Rio Jacui, as indicated in his former works on Rio 
Grande do Sul’s natural history (Ihering 1891, 1892). Therefore, 
Belton’s interpretation of Ihering’s citations is erroneous and clearly 
refers to the grassy hills around Herval (32°02'S, 53°24'W), a town 
located in the savanna domain (sensu Brasil 1986). Probably the 
distribution of C. cucullatus does not extend south of 31°40’S, 
coinciding with the limits of the originally forested region. 


RED-RUFFED FRUITCROW Pyroderus scutatus 


On 3 and 29 February 1995 we recorded two individuals at Arroio 
Cadeia and later found it regularly throughout the year at Arroio 
Andrade in numbers varying from one to five birds. Local residents 
informed us that in September and October groups of four or five 
individuals are seen in display, vocalizing intensely from the canopy. 
The nearest previous records and accepted southern limit for the 
species are Taquara (Ihering 1899) and Poco das Antas (Gliesch 1930), 
both located more than 250 km to the north of our study site. Besides 
these, Belton (1985) mentioned only two additional localities in Rio 
Grande do Sul with records of P. scutatus: Turvo state reserve and 
Garruchos. 


CHESTNUT-HEADED TANAGER Pyrrhocoma ruficeps 


We found this tanager only in four forest remnants: Arroio Andrade 
(up to three pairs), Arroio Cadeia (one pair), Arroio dos Porcos (three 
individuals) and Parque Farroupilha (male seen on 2 November 1993). 
Previously known in Rio Grande do Sul from Ihering’s (1899) record 
for Taquara, Gliesch’s (1930) for Santo Angelo and Camargo’s (1962) 
for Farroupilha. According to Belton (1985), the distribution of P. 
ruficeps covers forests north of 30°S and east of 54°W, and across the 
state north of 28°30’S (being absent from the highest areas of the 
extreme northeast), with a record from the southeastern hills near 
Santana da Boa Vista. Our records extend the known distribution of 
this tanager c. 100 km to the south. 


G. N. Mauricio & R. A. Dias 24 Bull. B.O.C. 1998 118(1) 


RUBY-CROWNED TANAGER Tachyphonus coronatus 


Recorded in all forest fragments studied, including Cerro das Almas. 
We also recorded it on two occasions at sea level, in swampy forests at 
Pontal da Barra marsh. These records extend the species’ known 
distribution by c. 250 km to the south, since it was previously known 
only from localities north of the central trough: Taquara (Ihering 
1899), Nova Wurtemburg (Pinto 1944), Osorio (Camargo 1962) and 
along the escarpment from Torres to the upper Rio Jacui, thence north 
to the Rio Uruguai and downstream to Garruchos (Belton 1985). 


VIOLACEOUS EUPHONIA Euphonia violacea 


On 28 February 1995 a male was seen in a mixed-species flock with 
Heliobletus contaminatus and Tangara preciosa at mid-level in the 
Arroio Cadeia forest. Previously known in Rio Grande do Sul from 
‘Taquara (Ihering 1899), Porto Alegre (Gliesch 1930, Camargo 1962) 
and Novo Hamburgo (Pinto 1944). Belton (1985) recorded it from the 
foot of the escarpment in the northeast and once near Garruchos. The 
nearest previous record is located c. 180 km to the north of our study 
site. 


GREEN-THROATED EUPHONIA Euphonia chalybea 


We first recorded a pair of this euphonia at Arroio dos Porcos on 
12 December 1994 and later saw three individuals (a male and two 
females) at Arroio Andrade on 28 May 1995. Mentioned from ‘Taquara 
by Ihering (1899), Poco das Antas by Gliesch (1930) and Novo 
Hamburgo and Nova Wurtenburg by Pinto (1944). According to 
Belton (1985) this species is uncommon near the southern escarpment 
and in scattered localities of the north-central region. Our records 
extend the known distribution of E. chalybea c. 200 km southwards. 


Acknowledgements 


We are deeply indebted to José Fernando Pacheco and Paulo Sérgio M. da Fonseca for 
providing material support, suggestions on the manuscript and constant encouragement. 
Dr Jurgen Haffer kindly provided valuable information and suggestions on the 
manuscript. William Belton and Mary LeCroy greatly helped in furnishing information 
on specimens at the American Museum of Natural History. We also thank Bret M. 
Whitney for his assistance in identifying some of the voice records and Morevy M. Cheffe 
for calling our attention to the precise location of Serra do Herval. Gabriel Viégas of Soft 
Shop Informatica patiently produced the map of the localities. We are extremely grateful 
to Nilson Loeck and the Rodeghiero and Lichnow families, whose hospitality ensured 
our fieldwork. 


References: 

Belton, W. 1984. Birds of Rio Grande do Sul, Brazil. Part 1. Rheidae through 
Furnariidae. Bull. Amer. Mus. Nat. Hist. 178: 371-631. 

Belton, W. 1985. Birds of Rio Grande do Sul, Brazil. Part 2. Formicariidae through 
Corvidae. Bull. Amer. Mus. Nat. Hist. 180: 3-241. 

Belton, W. 1994. Aves do Rio Grande do Sul: distribuigdo e biologia. Editora Unisinos, Sao 
Leopoldo. 

Berlepsch, H. v. & Ihering, H. v. 1885. Die V6gel der Umgegend von 'Taquara do Novo 
Mundo, Prov. do Rio Grande do Sul. Zeitschrift fiir dié gesammte Ornithologie 2: 
97-184. Tab. VI-IX. Budapest. 


F. G. Stiles 25 Bull. B.O.C. 1998 118(1) 


Brasil 1986. Projeto RADAMBRASIL, levantamento de recursos naturais. Vegetacdao. 
(Vol. 33, folhas SH 22, SH 21, SI 21—Porto Alegre, Uruguaiana, Lagoa Mirim). 
Rio de Janeiro. 

Camargo, O. R. 1962. Aves Sul-Riograndenses do Museu de Caga e Pesca. Pesquisas, 
Zool. 14: 1-67. 

Gliesch, R. 1930. Lista das aves colligidas e observadas no Estado do Rio Grande do Sul. 
Egatea 15: 276-292. 

Haffer, J. 1974. Avian speciation in tropical South America, with a systematic survey of 
the Toucans (Ramphastidae) and Jacamars (Galbulidae). Nuttall Orn. Cl., Publ. 14. 

Ihering, H. v. 1891. As arvores do Rio Grande do Sul. Annuadrio do Estado do Rio Grande 
do Sul para o Anno 1892: 164-196. 

Ihering, H. v. 1892. Os mamiferos do Rio Grande do Sul. Annuario do Estado do Rio 
Grande do Sul para o Anno 1893: 96-123. 

Ihering, H. v. 1899. As aves do Estado do Rio Grande do Sul. Annuario do Estado do Rio 
Grande do Sul para o Anno 1900: 113-154. 

Naumburg, E. M. B. 1935. Gazetteer and maps showing collecting stations visited by 
Emil Kaempfer in eastern Brazil and Paraguay. Bull. Amer. Mus. Nat. Hist. 68: 
449-469. 

Naumburg, E. M. B. 1937. Studies of birds from eastern Brazil and Paraguay, based on 
a collection made by Emil Kaempfer. Bull. Amer. Mus. Nat. Hist. 74: 139-205. 
Naumburg, E. M. B. 1939. Studies of birds from eastern Brazil and Paraguay, based on 
a collection made by Emil Kaempfer. Bull. Amer. Mus. Nat. Hist. 76: 231-276. 
Pinto, O. M. de O. 1938. Catalogo das aves do Brasil, 1* parte. Rev. Mus. Paulista 22: 

1-566. 

Pinto, O. M. de O. 1944. Catalogo das aves do Brasil, 2° parte. Departamento de 

Zoologia, Secretaria da Agricultura, Industria e Comércio, S. Paulo. 


Addresses: Giovanni N. Mauricio, Rua Gongalves Chaves 3448, 96015-560 Pelotas, RS, 
Brasil. Rafael A. Dias, Rua Celso Sellas 56, 96055-810 Pelotas, RS, Brasil. 


© British Ornithologists’ Club 1998 


Notes on the biology of two threatened 
species of Bangsia tanagers in northwestern 
Colombia 


by F. Gary Stiles 
Received 30 December 1996 


The genus Bangsia includes five species of chunky, thick-billed, rather 
short-tailed and strong-legged, medium-sized (body mass 35-45 g) 
tanagers. All of the species occur in very wet primary to lightly 
disturbed forests in upper tropical and subtropical life zones and all 
have restricted geographic distributions, making them potentially 
vulnerable to deforestation (Hilty 1985, Collar et al. 1992). The most 
threatened and least known species of the genus are undoubtedly the 
Black-and-Gold Tanager B. melanochlamys, and the Gold-ringed 
Tanager B. aureocincta (Isler & Isler 1987, Ridgely & Tudor 1989, 
Collar et al. 1992), both endemic to northwestern Colombia. 

B. melanochlamys had been recorded previously from two small areas 
some 150 km apart, on opposite sides of the Cauca valley: the north- 
western tip of the Central Andes between 1500 and 2385 m in the 
Department of Antioquia, and the southern and western slopes of 


F. G. Stiles 26 Bull. B.O.C. 1998 118(1) 


Cerro Tatama on the Pacific slope of the Western Andes between 1000 
and 2150 m, near the junction of the Departments of Valle del Cauca, 
Choco and Risaralda. In the former area, at least 25 specimens were 
collected between 1914 and 1948, but the region has since been severely 
deforested and there are no recent records; this population is feared to be 
extinct (Hilty 1985, Collar et al. 1992). There are a number of specimens 
and two recent (1987) records from the area of Cerro ‘Tatama: a speci- 
men taken at 1000 m, and a sighting of one individual (in several days of 
field work) at c. 1500 m (Collar et al. 1992, Pearman 1993), the latter 
providing the only information on the species in life. Available infor- 
mation on B. aureocincta was even more scanty: the species was known 
from just four specimens taken between 1909 and 1946 at three localities 
between 2040 and 2195 m in the same area of Cerro Tatama as the 
previous species; aside from the habitat notation of ‘““wet mossy forest”’ 
on a specimen label, nothing was known of its natural history. 

‘This paper presents field observations on the behaviour and ecology, 
measurements and weights of both species that greatly augment 
previous information. These data were obtained in the course of a 
biological inventory of northwestern Risaralda Department carried out 
between 1991 and 1993 by personnel of the Instituto de Ciencias 
Naturales through a contract with the Corporacion Autonoma Regional 
de Risaralda (CARDER); further data on the avifauna are given by 
Stiles (1992). Observations were made with 10 X 25 binoculars; birds 
captured in mist nets were measured with dial calipers or (wing chord, 
tail length) a millimeter rule (precision 0.1 or 0.5 mm, respectively; see 
Baldwin et al. 1931 for techniques) and weighed to the nearest 0.1 g 
with “‘Pesola”’ spring balances. 

Observations were made at the following sites: (1) Finca El 
Empalado, 9km N Mistrato: a large (>100 ha) patch of forest just 
northwest of the divide between the Pacific and Cauca Valley slopes of 
the Western Andes between c. 1600 and 1800 m, headwaters of the 
Quebrada Sutt; lightly high-graded forest on steeply sloping terrain, 
with abundant epiphytes and moss, frequent mist and high rainfall: 28 
March-3 April 1992. (2) Finca La Argentina, c. 4km SSW San 
Antonio del Chami and 3 km NW of El Empalado at a lower elevation 
(1250 m) in the drainage of the Quebrada Sutt; a smaller patch (c. 5 ha) 
of more heavily disturbed forest, connected to the previous site by a 
narrow strip of forest along the stream; observations by Sandra Arango, 
1-4 April 1992. (3) Alto de Pisones, a mountain ridge 16km E San 
Antonio del Chami and 9 km by trail NW Geguadas, a small village 
overlooking the Rio San Juan; observations were made in nearly 
pristine cloud forest between 1400 and c. 1850 m, 28 May—7 June 1992 
and 12-17 April 1993. A more detailed description of the vegetation is 
given by Salaman & Stiles (1996); the avifauna is treated in more detail 
by Stiles (1992 and in prep.). 


Bangsia melanochlamys 

At El Empalado, this species was fairly common (5-10 sightings 
daily) in forest canopy and along the forest edge on steep slopes. 
Individuals or presumed pairs frequently accompanied mixed flocks of 


F. G. Stiles zt Bull. B.O.C. 1998 118(1) 


tanagers (Tangara spp., Anisognathus spp., Chlorospingus semifuscus, 
Chlorochrysa phoenicotis) and other species in the canopy and sub- 
canopy, although I more often saw individuals or pairs by themselves, 
especially at fruiting Miconia trees. On one occasion, my attention was 
drawn to a bird hopping and calling excitedly with sharp tszp notes in a 
high, exposed treetop; it suddenly dived into cover an instant before 
a small hawk (Accipiter collaris) arrived at the same perch. On 30 March 
1992, I observed an individual, evidently a female, repeatedly carrying 
moss, fibres and leaf petioles into a mass of bromeliads.in the fork of a 
tree trunk 8 m above ground at the edge of a natural treefall clearing, at 
c. 1750 m. The presumed nest was hidden in the mass of epiphytes. A 
male was singing from an adjacent treetop while this individual worked. 
I captured three males with large cloacal protuberances (two collected, 
both with very enlarged testes) and a female with a well-developed brood 
patch at this site. At La Argentina, this species was observed twice, and 
one female with a brood patch was trapped and released. 

At Alto de Pisones, I observed B. melanochlamys frequently 
between 1400 and 1650 m and occasionally to 1750 m in May—June 
1992, nearly always in tall forest on steep slopes; a male sang 
regularly from a treetop at 1600m. Most birds seen occupied the 
middle levels to the lower canopy of the forest; about half of my 
observations were of birds accompanying mixed flocks (typically 
dominated by several species of tanagers, but containing a wider 
variety of other species than at El] Empalado), the others of birds 
resting or foraging alone or in pairs. I also noted two family groups 
(a pair with one or two full-grown, begging juveniles) in the upper 
understorey. In a net set along the ridge crest at 1730 m, I captured 
and collected a young male (testes very small, skull 10% ossified), 
probably recently independent, and measured and released an adult 
male and female. 

In April 1993, this species was common up to c. 1550m, and 
relatively scarce up to 1650-1680 m. Four males were noted singing 
from treetops along the mule trail between 1400 and 1550 m, one at 
1575 m, and one at 1650 m. Most individuals seen were in pairs, two of 
which were feeding more recently-fledged, stub-tailed juveniles (at 
1450 and 1550 m). At most a third of the individuals seen were with 
mixed flocks, which on the whole were less evident than during the 
1992 study period. 

Birds with flocks mostly foraged for insects, gleaning in moss tufts, 
epiphytes, and along small to medium-sized branches, sometimes 
leaning far forward to scan the undersides of branches and occasionally 
hanging upside-down to reach an insect, fruit, or flower. Birds foraging 
for fruit hopped heavily and rapidly along branches, plucking fruits 
while perched and usually crushing them in the bill, discarding husks 
and larger seeds in situ. Fruits taken included those of Cavendishia and 
Psammisia spp. (Ericaceae), Marcgravia sp. (Marcgraviaceae), Miconia 
and Topobaea (Melastomataceae), and those of an _ unidentified 
mistletoe, as well as the arillate seeds of Clusia. Flowers of Cavendishia 
were sometimes plucked, crushed at the base to extract the nectar, and 
discarded (see also Pearman 1993). Stomachs of collected birds 


F. G. Stiles 28 Bull. B.O.C. 1998 118(1) 


contained 75-100% fruit, the remainder insects; at least one of the 
fledglings seen in 1993 was being fed fruit (probably Marcgravia). 

The song of B.melanochlamys consists of 3—5 phrases that sound like 
pit-psEEyee or tst-tzEEeee, delivered rapidly and followed by a pause of 
several seconds before the next group. The tst note is staccato, the 
tzEEyee very high and penetrating, the first part with an explosive 
quality. The usual contact note is a sharp, staccato tst or pit; lone birds 
occasionally give a longer pseee or pseeyee, evidently the note described 
by Pearman (1993). 


Bangsia aureocincta 

I observed this species only at Alto de Pisones, where it was common 
to abundant above 1700m, progressively less common down to 
c. 1600 m (1993) or 1530 m (1992). Most birds were seen along the 
main ridge (Cuchilla de Gebania) running roughly E—W and separating 
the Rio Agtiita and Rio Mistrat6 watersheds. On 28 May 1992, along 
roughly 500 m of this ridge between c. 1730 and 1850 m, I counted at 
least 16 individuals including two family groups of a pair with one 
or two juveniles; at two points some 300 m apart, males were singing 
from the lower canopy. In April 1993, I noted four singing males 
near or along this ridge between 1700 and 1800 m, separated by 
distances of c. 100-250 m. A female was seen carrying moss into a mass 
of epiphytes on a thick horizontal limb some 15 m above the ground 
and c. 50 m below the ridge crest, but the nest was completely hidden. 
No family groups or fledglings were seen. In general, this species 
seemed less numerous in the area than during the previous year, 
possibly because its centre of distribution had shifted to higher 
elevations for breeding. 

Especially during the 1992 observations, this species was often 
associated with mixed flocks that included Tangara spp., Chloro- 
thraupis stolzmanni, Chlorospingus flavigularis and/or semifuscus, 
Chlorochrysa phoenicotis, Euphonia xanthogastra as well as furnariids, 
woodcreepers, antwrens, barbets and other species; however, it rarely 
occurred in the same flocks as B. melanochlamys, and even at fruiting 
Miconia trees (common along the trail to Puerto de Oro above 
c. 1650 m) the two species seldom coincided in their visits (and were 
not observed to interact when they did so). B. aureocincta occurred on 
average slightly lower in the vegetation than melanochlamys, typically 
from the upper understorey through the midcanopy. With flocks, its 
foraging was mainly for insects and similar to that of melanochlamys: 
searching deliberately in moss and epiphytes, mostly along rather thick, 
horizontal branches. Individuals or pairs were also recorded taking 
fruits of Cavendishia, Marcgravia, Guettarda (Rubiaceae), an un- 
identified mistletoe and Anthurium (Araceae), and arillate seeds of 
Clusia and ?Tovomita (Guttiferae). 

Between 29 May and 2 June 1992 I captured 7 individuals in mist 
nets set along the ridge at 1730-1750 m. An adult male and female 
and a subadult male were collected; two males and two females were 
measured and released. All individuals except one female were 
commencing the annual moult; the males collected had enlarged testes 


F.G. Stiles 29 Bull. B.O.C. 1998 118(1) 


TABLE 1 
Measurements (mean, standard deviation, range) of Bangsia tanagers captured in NW 
Risaralda Department, Colombia, 1992-1993 


B. aureocincta B. melanochlamys 


Males (n=4) Females (n=3) Males (m=5) Females (n=3) 
Exposed culmen 14.03, 0.46 13.70, 0.26 13.47, 0.48 13.07, 0.16 
(13.6-14.6) (13.5-14.0) (12.9-14.0) (12.9-13.2) 
Total culmen 18.10, 0.60 17.97, 0.90 17.55, 0.42 1722330.25 
(17.2-18.4) (17.3-19.0) (17.4-18.0) (17.0-17.5) 
Commissure width 13.80, 0.91 13.83, 0.42 11.96, 0.39 11.96, 0.25 
(12.6—-14.6) (13.5-14.3) (11.6-12.6) (11.7-12.2) 
Bill depth at nostril 8.15, 0.19 8.10, 0.26 7275;30i13 7.87, 0.15 
(7.9-8.3) (7.9-8.4) (7.5-8.0) (7.7-8.0) 
Tarsus length 26.08, 0.91 FAs tes hie See 4!) 24.72, 0.94 24.60, 0.36 
(24.8-26.5) (24.7-27.4) (23.8-25.8) (24.3-25.0) 
Wing chord 89.40, 1.04 84.19, 1.05 So. 50, Tord 5 ypu) Js ha | 
(88.0—90.5) (83.0-85.0) (84.0-—86.5) (80.5-83.5) 
Tail length 52.42, 0.68 52.23.0275 45.06, 0.88 44.96, 1.50 
(51.5-53.5) (51.5-53.0) (44.0-47.0) (43.5-46.5) 
Body mass (g) 42.05, 3.40 Ce Os) 38.80, 0.81 35.65, 0.92 
(39.4-46.6) (38.3-44.6) (37.7-39.4) (35.0-36.3) 


(9x 8mm and 64mm for the adult and subadult, respectively), 
while the females had old brood patches and the collected female had a 
postreproductive ovary. Stomachs of collected birds contained 70—90% 
fruit, mainly Muiconia, Cavendishia and Marcgravia, the remainder 
insects. 

The song of B. aureocincta consists of sharp, penetrating, high- 
pitched whistles or thin, watery trills tseewurr delivered in groups of 
3-6. The individual whistles start explosively and slur downwards; 
usually the first one or two are shorter than the rest. A frequently given 
note in situations of alarm or excitement is a short, twittering trill on a 
lower pitch; when moving together, individuals may maintain contact 
with sharp chip or chit notes, heavier than the corresponding notes of 
B. melanochlamys. 


Measurements, weights and plumages 

B. aureocincta is slightly larger in all measurements, but apart from 
this the two species are quite similar in overall proportions. In both 
species males have significantly longer wings than females (Mann-— 
Whitney U-tests), but the sexes do not differ in any other measurement 
(save that males of B. melanochlamys weighed more than females, which 
may be an artefact of the small sample size; see Table 1). 

Sexual differences in plumage in B. melanochlamys are too slight to 
be evident in the field: the head and back of the female are a slightly 
duller, less glossy black and the orange of the breast is loss glossy 
and intense. The sexes differ much more in B. aureocincta and, given 
the uncertainty regarding the adult female plumage (Isler & Isler 1987) 


F. G. Stiles 30 Bull. B.O.C. 1998 118(1) 


due to the scanty data of the few available specimens, I here describe 
the plumages of my specimens in some detail. ADULT MALE: 
head, throat, sides of breast glossy black; chin whitish, scaled with 
dull black; postocular and malar stripes and postauricular bar bright 
yellow, forming a ring around the cheeks and auriculars; centre of 
breast bright orange-yellow, remaining underparts bright olive-green, 
washed with yellow medially and on crissum. Back dark moss-green, 
rump and upper tail-coverts paler, brighter green. Central rectrices 
duller, darker olive; remaining rectrices dusky, edged with olive, faintly 
tinged with dull blue; wings blackish, the wing-coverts and secondaries 
broadly edged with rather dull, dark blue. Iris dark red; maxilla black, 
mandible yellowish horn colour; tarsi and feet light greyish-brown. 
ADULT FEMALE: crown and nape dull blackish smudged with dull 
dark blue; auriculars and cheeks dark moss green, the yellow ring 
surrounding them narrower and more greenish-yellow; chin yellowish- 
white smudged with dusky, throat dusky-olive shading to olive-green on 
the sides of the breast; yellow of breast duller, less orange; posterior 
underparts olive green with less yellow wash medially; green of dorsum 
duller, more olive; blue of wings duller. Mandible mostly chrome 
yellow, tip brownish; soft parts otherwise like male. SUBADULT 
MALE (probably one year old, as skull completely ossified): black of 
head dull, not glossy, slightly smudged with dull, dark blue on crown 
and nape; auriculars and sides of breast dull, dark greenish-black; 
pale area of chin tinged yellowish and more extensive than in adult; 
yellow of breast duller, smudged with dusky; back slightly paler and 
duller; iris dark chestnut; blue of remiges slightly greyer; otherwise like 
adult male. 


Discussion 

Breeding of both species appeared to be at its height during March 
and April, with many well-grown young in late May-—early June. There 
appeared to be a fairly well-marked separation in elevation during 
breeding at Pisones, with melanochlamys mostly below 1600-1650 m 
and aureocincta mostly 1700m and higher; more overlap evidently 
occurred following breeding. At E] Empalado, where aureocincta was 
not observed, melanochlamys nested up to at least 1750 m. A difference 
in preferred habitat may also occur, with melanochlamys occupying 
steeply sloping terrain, aureocincta the ridge crests. 

In plumage pattern including the relative lack of sexual dimorphism, 
vocalizations and behaviour, melanochlamys seemed quite similar to 
B. arcae of Central America (cf. Isler & Isler 1987, Stiles & Skutch 
1989, Ridgely & Gwynne 1989); the two are doubtless closely related, 
and probably comprise a superspecies. The closest relative of B. 
aureocincta is probably B. edwardsi to the south of the Rio San Juan 
drainage (Hilty & Brown 1986, Ridgely & Tudor 1989); only from this 
species have I heard a similar, trilling alarm note. However, the two 
differ much more in plumage pattern, and aureocincta shows. much 
greater sexual dimorphism. 

The Alto de Pisones region of western Risaralda would appear to be a 
critical conservation area for both species, especially for B. auwreocincta 


F. G. Stiles 31 Bull. B.O.C. 1998 118(1) 


which at present is known to exist nowhere else. CARDER is preparing 
to execute a management plan for this area, which may also be included 
in the proposed Caramanta National Park. However, because the area 
is also part of an Embera Indian reservation and recently has been 
invaded by guerrilla groups, the situation is complicated to say the 
least. Potential threats to the area resulting from the construction of the 
Geguadas—Santa Cecilia road were outlined by Salaman & Stiles 
(1996). It is hoped that funds deriving from the Species Sponsorship 
Program of BirdLife International (Alto de Pisones is the type locality 
of the recently described Vireo masteri; Salaman & Stiles 1996) will 
help CARDER to act in spite of the difficult situation. It will also be 
important to survey other mountain ridges around the massif of Cerro 
Caramanta to determine whether other populations of B. aureocincta 
exist in this region. 


Acknowledgements 


I thank Gonzalo Andrade, Diana Mora, Milton Munoz and Pablo Rodriguez of the 
Universidad Nacional de Colombia, and Carlos Alberto Vanegas of CARDER, for help 
in the field. Sandra Arango provided observations from La Argentina. Logistics were 


provided through the Instituto de Ciencias Naturales, and facilitated by Eduardo 
Londono of CARDER. 


References: 

Baldwin, S. C., Oberholser, H. C. & Worley, L. G. 1931. Measurements of Birds. Sci. 
Publ. Cleveland Mus. Nat. Hist., no. 2. 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
T. A. & Wege, D. C. 1992. Threatened Birds of the Americas: the ICBP/IUCN Red 
Data Book (3rd edn, part 2). ICBP, Cambridge, U.K. 

Hilty, S. L. 1985. Distributional changes in the Colombian avifauna: a preliminary blue 
list. Pp. 1000-1012 zm P. A. Buckley et al. (eds), Neotropical Ornithology. Orn. 
Monogr. no. 36. 

Hilty, S. L. & Brown, W. L. 1986. A Guide to the Birds of Colombia. Princeton Univ. 
Press. 

Isler, M. L. & Isler, P. R. 1987. The Tanagers: natural history, distribution and 
identification. Smithson. Inst. Press. 

Pearman, M. 1993. Some range extensions and five species new to Colombia, with notes 
on some scarce or little known species. Bull. Brit. Orn. Cl. 113: 66-75. 

Ridgely, R. S. & Gwynne, J. A. 1989. A Guide to the Birds of Panama including Costa 
Rica, Nicaragua and Honduras. Princeton Univ. Press. 

Ridgely, R. S. & Tudor, G. 1989. The Birds of South America. Vol. 1. Univ. Texas Press. 

Salaman, P. G. W. & Stiles, F. G. 1996. A distinctive new species of Vireo 
(Passeriformes, Vireonidae) from the Western Andes of Colombia. Ibis 138: 
620-629. 

Stiles, F. G. 1992. Un inventario preliminar de la avifauna del noroccidente del 
Departamento de Risaralda. Report to CARDER, Inst. Cienc. Naturales. 

Stiles, F. G. & Skutch, A. F. 1989. A Guide to the Birds of Costa Rica. Cornell Univ. 
Press. 


Address: F. Gary Stiles, Instituto de Ciencias Naturales, Universidad Nacional de 


Colombia, Apartado 7495, Santafe de Bogota, Colombia. email: fstiles@ciencias. 
ciencias.unal.edu.co 


© British Ornithologists’ Club 1998 


E. M. Figueroa-Esquivel et al. 32 Bull. B.O.C. 1998 118(1) 


New distributional information on the birds of 
southern Quintana Roo, México 


by Elsa M. Figueroa-Esquivel, Adolfo G. Navarro S. & 
Carmen Pozo-de la Tijera 


Received 27 Fanuary 1997 


The avifauna of the Mexican state of Quintana Roo, located in the 
eastern half of the Yucatan Peninsula, has received much attention 
recently, partly because of its importance as a wintering ground for 
many Neotropical migrants (Scott et al. 1985, Gatz et al. 1985; 
Chavez-Léon 1988, Lopez-Ornat 1989). Since the pioneering 
monograph of Paynter (1955), the Yucatan Peninsula has been 
recognised as an important area of avian diversity and endemism, 
particularly the arid northern section. However, knowledge of the 
avifauna of Quintana Roo has been concentrated in the northern part 
and on Cozumel Island, and much of the south remains poorly known 
(Paynter 1955, MacKinnon 1992, Vasquez et al. 1992, Figueroa 1994). 

The Museo de Zoologia of the Colegio de la Frontera Sur, Chetumal 
(ECOSUR, formerly Centro de Investigaciones de Quintana Roo, 
CIQRO) surveyed the vertebrate fauna in the poorly known southern 
section of the state between August 1992 and August 1993. Field work 
was conducted in seven localities within the Municipio of Othon 
Pompeyo Blanco in southernmost Quintan Roo (17°53’-18°13'N, 
88°46'-89°15'W). The elevational range of the area is from 0 to 200 m. 
Dominant vegetation throughout is tropical semideciduous forest (sensu 
Rzedowski 1988), with dominant trees including Bursera simaruba, 
Manilkara zapota, Brosimum alicastrum, Metopium browne, and Chryso- 
phila argentea (Torres, 1991). Large tracts of original vegetation have 
been modified due to human disturbance, mainly for cattle grazing, 
agriculture, and exploitation of fine hardwoods. General coordinates and 
elevation of localities mentioned in the species accounts are as follows: 
4:-7km N, 13 km W of Calderon (18°07'N, 88°55’W; 200 m); Estero 
Franco (17°56’N, 88°52’W; 30 m); La Union (17°56'N, 88°51'W; 0 m); 
Los Tornillos (18°05’N, 89°03’W; 130m); Dos Aguadas (18°07'N, 
89°08'W; 180m); 2.3km S of Nuevo Veracruz (18°02'N, 89°10'W; 
140 m); El Naranjal (18°13'N, 89°02'W; 140 m). 

During 65 days of field work, mist-netting and sight or auditory 
records were used to inventory the species present in the area: We 
collected selected specimens of as many species as possible; voucher 
specimens are at the Museo de Zoologia, ECOSUR (MZECOSUR) 
and at the Museo de Zoologia, Facultad de Ciencias, Universidad 
Nacional Aut6noma de México (MZFC). We report here seven species 
apparently new or noteworthy for the state. 


MISSISSIPPI KITE [ctinia mississippiensis 
On 20 October 1992, an individual was observed at Los Tornillos. 
This species is a transient along the east coast and southeastern Mexico 


E. M. Figueroa-Esquivel et al. 33 Bull. B.O.C. 1998 118(1) 


to South America. No previous records exist from the Yucatan 
Peninsula, except for one in Yucatan (no locality given, Howell & 
Webb 1995). Possibly overlooked, and more widespread, this record is 
the first from Quintana Roo. 


SWALLOW-TAILED KITE Elanoides forficatus 

Six individuals were observed on 12 May, 1993, at Los Tornillos. In 
June 1994, a nest was discovered on a dead tree on the edge of a 
cornfield at Isidro Fabela, central Quintana Roo (De Alba 1997); in 
June 1995, an individual was observed 5km S Dos Lagunas, in 
southeastern Campeche (Figueroa & Salgado in prep.). This species 
is considered a common winter transient throughout the Yucatan 
Peninsula, and a local breeder in eastern Mexico (eastern Chiapas) and 
Central America (Rappole et al. 1993, Howell & Webb 1995). The nest 
record extends the Mexican breeding range significantly northwards. 


WHITE-NECKED JACOBIN Florisuga mellivora 


On 23 March 1993, an adult male (heavy fat deposits, no moult) was 
collected 4-7 km N, 13 km W of Calderon (MZECOSUR A-512) in 
cultivated grassland approximately 1m high. The next day, two 
additional individuals were observed at the same locality, foraging at 
several species of flowers. On 14 May 1993, another was observed near 
Los Tornillos, in a patch of well preserved forest. ‘This species is 
known from rain forests in 'l‘abasco, Veracruz, northern Oaxaca and 
Chiapas (Miller et al. 1957, AOU 1983, Binford 1989, Howell & Webb 
1995) and adjacent northern Guatemala in the Petén (Land 1970) 
and Tikal (Smithe 1966), where it is considered uncommon. Records 
from Belize (Russell 1964) are from southernmost localities. No 
previous records exist from the Yucatan Peninsula in Mexico except for 
an observation from Calakmul Biosphere Reserve, in southeastern 
Campeche (PRONATURA 1993). 


BUFF-THROATED FOLIAGE-GLEANER Automolus ochrolaemus 


In April 1992, an adult male was collected 7km N of La Union 
(MZFC 10551). On 24 and 25 February 1993, a male and a female 
(MZECOSUR A-380, A-387) were collected 2.3km S of Nuevo 
Veracruz, in a well preserved patch of forest. The male showed no 
evidence of fat or gonadal enlargement; the female, however, had an 
enlarged ovary and light fat. These records are the first for the State; 
previous Mexican records near the study area are from Oaxaca, 
Tabasco and Chiapas in Mexico (AOU 1983). The species is 
considered uncommon in northern Guatemala (Land 1970), and rare in 
northern Belize (Russell 1964). 


THICK-BILLED SEEDFINCH Oryzoborus funereus 

Six specimens of this species (MZECOSUR A-157, A-538, A-610, 
A-612, A-613, A-641) were collected on 29 August 1992, and 25 March 
and 2, 3 and 24 June 1993. These are the first records for the state; 
previous peninsular records are those of Robbins, who sighted one 
individual at Rancho Santa Anita, Campeche (MacKinnon 1992). The 


E. M. Figueroa-Esquivel et al. 34 Bull. B.O.C. 1998 118(1) 


species has been recorded from Guatemala and Belize (Land 1970; 
Russell 1964), but seems to be very local in occurrence, as in Mexico 


(Howell & Webb 1995). 


FUERTES’ ORIOLE Icterus (spurius) fuertest 


One adult in male plumage was observed on 25 February 1993, 
2.3km S of Nuevo Veracruz among a flock of Orchard Orioles Icterus 
(spurius) spurius. It was not possible to determine if females or 
immatures of this species were also present. The specific status of the 
two forms is still in debate, but winter sympatry has been documented 
elsewhere (e.g. AOU 1983). This record is the first for Quintana Roo. 


EASTERN MEADOWLARK Sturnella magna 


One individual was sighted on 4 June 1993, in a patch of grass at the 
margins of Rio Hondo. No previous records exists from Quintana Roo, 
and its seasonal status is uncertain; the data suggests a resident 
population (Russell 1964). This species is uncommon in Belize (Russell 
1964) and northern Guatemala (Land 1970). In the Yucatan Peninsula 
it has been recorded only along the northern coast of the state of 
Yucatan. 


Discussion 


Due to the lack of thorough surveys in many regions, knowledge of the 
Mexican avifauna is still fragmentary. Although organized surveys have 
been developed for several states and regions (e.g. Binford 1989, 
Navarro et al. 1993), many areas remain little known. The avifauna 
of southern Quintana Roo is very rich, and an analysis of general 
distribution patterns of bird species will be published elsewhere 
(Figueroa in prep.). Undoubtedly, most of the species still to be 
recorded have been overlooked due to lack of thorough field work, but 
some crop-associated species may well be recently established, as a 
result of the extensive transformation of forest to agriculture and 
pasture lands. Examples may include the seedfinch and meadowlark 
recorded here. 


Acknowledgements 


We would like to thank the personnel of the Museo de Zoologia, Colegio de la Frontera 
Sur (ECOSUR), especially Enrique Escobedo, Alejandro de Alba, Noemi Salas, Maximo 
Suarez, and Lazaro and Cirilo Rodriguez, for support and companionship in the field. 
Thanks to A. Townsend Peterson, David W. Snow, Juan Francisco Ornelas and Javier 
Salgado, who provided helpful suggestions and criticism in the preparation of this note. 
Financial support was obtained from ECOSUR. This paper constitutes the number 13 in 
the series ““‘New distributional information on Mexican birds’’. 


References: 

American Ornithologists’ Union. 1983. Check-list of North American Birds. 6th edn. 
A.O.U., Washington, D.C. 

Binford, L. C. 1989. A distributional survey of the birds of Mexican state of Oaxaca. Orn. 
Monogr. no. 43, American Ornithologists’ Union. 


E. M. Figueroa-Esquivel et al. 35 Bull. B.O.C. 1998 118(1) 


Chavez-Leon, G. 1988. Aves de Quintana Roo, México. SARH. Rev. Cien. For. 13(63): 
97-154. 

De Alba, A. 1997. Avifauna de la zona noroeste del Municipio Othon Pompeyo Blanco, 
Quintana Roo, México. Tesis Profesional, Universidad Nacional Autonoma de 
Mexico. 

Figueroa-Esquivel, E. M. 1994. Estudio avifaunistico de la region Sur del estado de 
Quintana Roo, México. Tesis Profesional, Universidad Nacional Autonoma de 
Mexico. 

Gatz, T., Gent, P., Jakla, M., Otto, R. & Ellis, B. 1985. Spotted Rail, Brant and 
Yellow-breasted Crake records from the Yucatan. American Birds 39: 871-872. 

- Howell, S. N. & Webb, S. 1995. A Guide to the Birds of Mexico and Northern Central 
America. Oxford Univ. Press. 

Land, H. C. 1970. Birds of Guatemala. Livingston Publishing ani 

Lopez-Ornat, A. 1989. New and noteworthy records of birds from the eastern Yucatan 
Peninsula. Wilson Bull. 101: 390-409. 

Mackinnon, B. (ed.) 1992. Check-list of the birds of the Yucatan Peninsula. Amigos de 
Sian Ka’an A.C. 

Miller, A. H., Friedmann, H., Griscom, L. & Moore, R. T. 1957. Distributional 
check-list of the birds of Mexico. Part II. Pacif. Coast Avif. 32. 

Navarro-Sigtienza, A. G., Hernandez Banos, B. & Benitez-Diaz, H. 1993. Las aves del 
estado de Querétaro, Mexico. List. Faun. Méx. Inst. Biol. UNAM 4. 

Paynter, R. A., Jr. 1955. Ornithogeography of the Yucatan Peninsula. Peabody Mus. 
Nail. Hist. Publ.. 9. 

PRONATURA Peninsula de Yucatan, A. C. 1993. At Calakmul: The Birds. Reporte de 
una especie nueva para la Peninsula de Yucatan, Florisuga mellivora, observada en la 
Reserva de la Biosfera de Calakmul, Campeche. Kambul. 3(2): 4. 

Rappole, J. H., Morton, E. S., Lovejoy, T. E. & Roos, J. L. 1993. Aves migratorias 
Nearticas en los Neotroépicos. Conservation and Research Center, National Zoological 
Park, Smithsonian Institution, Washington. 

Russell, S. M. 1964. A distributional study of the birds of British Honduras. Orn. 
Monogr. no. 1, American Ornithologists’ Union. 

Rzedowski, J. 1988. Vegetacion de México. Ed. Limusa, Mexico. 

SCT (Secretaria de Comunicaciones y Transportes) 1987. Mapa de Carreteras, Atlas 
Cultural de México. Cartografico II. SEP-INAH, Planeta, México. 

Smithe, F. B. 1966. The birds of Tikal. American Museum of Natural History. 

Scott, P. E., Andrews, D. & Mackinnon, B. 1985. Spotted Rail: first record from the 
Yucatan Peninsula, Mexico. American Birds 39: 854. 

Torres, P., S. A. 1991. Estudio Floristico de las Pteridofitas del Sur. del estado de 
Quintana Roo. Tesis Professional, Instituto Tecnologico de Chetumal. Chetumal, 
Quintana Roo, México. 

Vasquez, M. A., Munoz, A., March, I., Dominguez, R., Garcia, G., Garcia, J., Salazar, 
S. & Ordonez, M. 1992. Zona Maya: Un anialisis de las areas silvestres. Centro de 
Estudios para la Conservacion de los Recursos Naturales A.C. y Sostenibilidad 
Maya. San Cristobal de las Casas, Chiapas. México. 


Addresses: Elsa M. Figueroa-Esquivel, Museo de Zoologia, Colegio de la Frontera Sur 
Unidad Chetumal, ECOSUR. Apdo. Postal 424, Chetumal, Q. Roo 700. México. 
Present Address: ECOMAT, Universidad Aut6noma de Campeche, Av. Agustin 
Melgar s/n, Campeche, Campeche 24030, México. Adolfo G. Navarro-S., Museo de 
Zoologia, Facultad de Ciencias, Universidad Nacional Autonoma de México, Apdo. 
Postal 70-399, México, D.F. 04510, México. Carmen Pozo-de la Tijera, Museo de 
Zoologia, Colegio de la Frontera Sur Unidad Chetumal, ECOSUR, Apdo. Postal 
424, Chetumal, Q. Roo 7700, México. 


© British Ornithologists’ Club 1998 


R.C. Brace & }. Hornbuckle 36 Bull. B.O.C. 1998 118(1) 


Distributional records of and identification 
notes on birds of the Beni Biological Station, 
Beni, Bolivia 


by Robin C. Brace & Jon Hornbuckle 
Received 19 February 1997 


Distributional data on the Bolivian avifauna have accumulated rapidly 
in recent years, which has facilitated greatly the identification of 
localities crucial for conserving species at risk from extinction. Wege & 
Long (1995) identified 27 ‘Key Areas’, of which nine were ranked as 
being of top priority with regard to the numbers of threatened species 
occurring. One of these locations is the Beni Biological Station (BBS) 
(Estacion Biologica del Beni) (IUCN category I; IUCN 1992), which 
was the first ever UNESCO ‘Man and Biosphere’ site. This 160 000 ha 
tropical lowland reserve is characterized by a complex habitat mosaic, 
spanning rainforest, savanna and wetland; consequently it has a rich 
avifauna. A recent inventory has been provided by Brace et al. (1997), 
who list 478 species, and treat in detail the status of the four threatened 
and 15 near-threatened species (Collar et al. 1994) which had been 
recorded as of 1995. However, information on either range extensions 
or new observations relating to the identification of non-threatened 
species was not provided in that paper, an omission which is rectified 
here. 

The great majority of the data presented are derived from 
observations made during three annual (1994—96), 6-week visits to the 
BBS over the July-September period (dry season), accompanied by 
EarthCorps (Earthwatch) teams and Bolivian students, to undertake 
ecological research on the faunas of savanna-based forest islands (R. C. 
Brace et al., unpubl. data). However, commentaries on three of the 22 
species discussed are based solely on sightings made by White et al. 
(1993) in 1992. Earlier endeavours at the BBS by Cabot et al. (1986), 
Flores (1988), S. L. Hilty (unpubl. data), Rocha (1988, 1990) and other 
workers led to the production of a preliminary species catalogue by 
Miranda et al. (1991), which tabulated more than 400 species. Although 
that inventory acted as a valuable template for our expanded listing 
(Brace et al., loc cit.), 1t was necessary nevertheless to evaluate critically 
a number of records, which resulted in the exclusion of no less than 10 
species. 

Although we now have a good working knowledge of the avifauna of 
some parts of the BBS—notably those within easy reach of the El 
Porvenir field station located on the southern flank of the reserve— 
there is a paucity of information concerning centrally located tracts 
which are difficult to access. It is our hope that this paper will catalyse 
investigations of hitherto relatively unexplored areas of the BBS, and 
will encourage further study during the austral summer (wet season), a 
period over which there is little information, for example, on the influx 
of northern migrants. 


R.C. Brace & }. Hornbuckle ve Bull. B.O.C. 1998 118(1) 


BENI 
BIOL AL 


TATION (BB 
Llanos de Mojos 


PERU 


. 
4 


BRAZIL 


ath SANTA CRUZ 
a=? 7 Ung € 4 @ 
- Santa Cruz 


PARAGUAY 


Figure 1. Location of the Beni Biological Station (BBS). The administrative departments 
of Bolivia, and some cities and towns, are indicated, as are adjacent countries. 


Study location and areas 


The BBS is located in the Llanos de Mojos (Fig. 1), a lowland (c. 200 m 
altitude) plain characterized by savannas and forested areas. The 
savannas are either hyperseasonal (subject to alternating soil saturation 
and drought/fires) or seasonal (subject to an extended dry period) 
(Sarmiento 1983). The reserve itself extends over an area roughly 80 by 
30 km in extent, 70% of which is covered by a variety of forest types, 
though humid seasonal categories dominate (Miranda et al. 1991). The 
northern and southern limits of the reserve are demarcated, 
respectively, by the Rios Manique and Curiraba (Fig. 2a); inundation 
of the former, a ‘white-water’ river, is responsible for the presence of 
much swamp forest within the reserve core. The El] Porvenir (PVR) 
field station (14°52’S, 66°20’W), where we were based, is located 
180 km west of Trinidad and 50km east of San Borja. Planned 
expansion (Miranda pers. comm.) of the BBS will engulf the 2500 ha 
PVR estancia (Fig. 2b), which extends northwards to the Rio Curiraba. 


R.C. Brace & }. Hornbuckle 38 Bull. B.O.C. 1998 118(1) 


(a) N__ Rio Manique Laguna La Botella 


PSC 


Rio Aguas Negras 
San Borja Se Ae 
> Bor iti 7 


Laguna FNC 


Piraquinas 


Rio Aguas Negras 


TRC 
TRF estancia 


forest islands 


PVR estancia 
Road (To Trinidad) 


PVR field station 


Figure 2. (a) Map of the Beni Biological Station to show study locations and areas (see 
text): based on maps given in Miranda et al. (1991). To the southwest and northeast, the 
limits of the reserve (as of 1996) are indicated by dashed lines. The northern and 
southern limits are demarcated by the Rios Manique and Curiraba (with accompanying 
riverine forest shown) respectively. South of the latter les the El Porvenir (PVR) 
estancia. Stippled areas represent forest; unmarked tracts, in the east and adjacent to the 
San Borja—Trinidad road, designate savanna. To the east of PRV a representative 
selection of savanna-based forest islands/fragments is depicted. (b) Map showing the E] 
Porvenir estancia and affiliated field station, in more detail; some forest islands are 
depicted. Also denoted are the TRF estancia, the FFF, and some features south of the 


road. 


Consequently, this ornithologically well-worked estancia is, and has 
been considered (Brace et al. 1997), an integral part of the reserve. The 
same is true of the Triunfo (TRF) estancia, through which it is 
necessary to pass in order to reach forest camps from PVR. It should be 
noted that the reserve forest block is virtually isolated now from all 
adjacent forested areas by intervening savanna, giving concern in the 


R.C. Brace & }. Hornbuckle 39 Bull. B.O.C. 1998 118(1) 


context of putative future avifaunal impoverishment. However, 
negating this isolation to some degree is the presence of well-vegetated, 
former river beds (curiches) meandering across the PVR savanna, and of 
forest islands (islas des bosques), which appear to act, collectively, as 
habitat corridors (Saunders & Hobbs 1991). They are utilized by a wide 
selection of forest birds, though by no means all such species (R. C. 
Brace & J. H. Pearce-Higgins unpubl. data). 

The following areas are distinguished in the text; their locations are 
indicated in Figure 2. 


PVR estancia—savanna. This fairly open terrain is dominated by 
Andropogon bicornis (Graminaceae) (up to c. 80cm in height), and 
contains scattered fire-resistant trees: e.g. Tabebuia aurea, T. chrysantha) 
(tajibos) (Bignoniaceae), Pseudobombax spp. (Bombacaceae). It is 
subjected to limited seasonal inundation, cattle grazing, and to annual 
incursions of fire started in neighbouring estancias during the 
August/September period to promote the growth of new grass. The 
fires frequently affect quite extensive areas; some controlled, ‘in-house’ 
burns are initiated also. Consequently, the savanna exhibits at any one 
time a patchwork of grass of different heights. 

PVR _ estancia—forest islands. Dotted in the PVR savanna are 
numerous such islands ranging in size from <1 to 5+ ha. Guazuma 
ulmifolia (coco) (Sterculiaceae), Ficus spp. Curatella americana 
(Moraceae) and Scheelea princeps (motact) (Palmaceae) are typical 
island trees; fire-resistant species predominate on the periphery. The 
centres of most islands are relatively devoid of undergrowth due to 
cattle grazing, but narrow foliated scrubby borders persist which attract 
a wide variety of birds. 

Florida estancia—forest fragment (FFF). An 11 ha fragment created 
some 20 years ago as a result of road construction. Although grazed, the 
western portion retains fairly dense undergrowth and the periphery is 
profusely vegetated. 

Laguna Normandia (LGN )—cyperacean edge. A fringe (up to 20 m 
wide) of Cyperus giganteus (Cyperaceae) (to 2 m in height) punctuated 
in places by other sedges, grasses and some bushes. 

Triunfo (TRF) estancia. An area of savanna which is somewhat less 
open than that of the PVR estancia, with some patches of chaco-like 
scrub. 

El Trapiche (TRC). A camp sited in low (<15m), seasonally 
inundated palm forest, 700 m distant from the savanna edge and 250 m 
north of the Rio Curiraba. Mist-netting locations were up to 1 km 
north of the camp, and within a narrow belt of riverine forest on the 
southern flank of the Curiraba, where there are many tangled bushes. 

Pascana (PSC). A camp set in swamp forest alongside a small 
‘black-water’ lagoon; situated 6 km north of Trapiche. 

08 (Zero Ocho). A Chimane Indian village sited adjacent to the 
Rio Curiraba. Mist-nets were set up in seasonally inundated forest 
c. 0.5-1 km northwest of the village, accessed by a track leading to PSC. 

Final Camp (FNC). A location in the northeast of the reserve 
alongside the Rio Manique. The camp was located in high (to 30 m) 


R.C. Brace & }. Hornbuckle 40 Bull. B.O.C. 1998 118(1) 


riverine and swamp forest, which is quite open in places (White et al. 
1993). 


Systematic ordering in the species accounts is conservative and 
follows Clements (1991). For the tyrant flycatchers, the taxonomy and 
English names used adhere closely to Ridgely & Tudor (1994). Species 
new for the BBS (1992-96) are indicated by an asterisk; species 
additional to those listed by Brace et al. (1997) are denoted by two 
asterisks (four species—taking the reserve total to 482). In those 
instances where only one of the authors observed a species, the 
appropriate initials appear; ” signifies that neither author was involved 
in the sighting concerned (two species in 1992). JWP-H refers to James 
Pearce-Higgins who accompanied us in 1995, and BMNH to the 
Natural History Museum, Tring. 


Species accounts 
PEARL KITE Gampsonyx swainsonu* 


One watched in savanna 3km north of PVR on 17 September 1992 
(RCB et al.) (White et al. 1993) is the first record for Beni; it remains 
the sole sighting for the BBS. This species was known previously from 


the non-Amazonian lowlands of Santa Cruz and Taria, and from La 
Paz (Parker 1989). 


TINY HAWK Accipiter superciliosus** 

An individual seen near TRC on 29 July 1996 (George and Joan 
Hardie, RCB) is the first and only record for the BBS, although both a 
pair (28 August 1994) and a single bird (JH) (August 1995) have been 
observed in secondary forest with clearings, only 4 km to the south of 
the PVR estancia. These sightings would appear to be the second to 
fourth reports for Beni, the first coming from foothill forest (c. 600 m) 
40 km west of San Borja in 1990 (Parker 1989, Parker et al. 1991). 
Within Bolivia, the species is known only from Beni, Cochabamba and 
Santa Cruz (Remsen & Traylor 1983, Arribas et al. 1995). 


BURROWING OWL Speotyto cunicularia* 


A pair in residence at El Porvenir in 1995 and 1996 constitute, 
surprisingly, the first reports from the department of Beni. 


LITTLE NIGHTJAR Caprimulgus parvulus* 

Although widespread in lowland Bolivia (Arribas et al. 1995), this 
species had not been recorded at the BBS until August 1994, when it 
was heard in the savanna adjacent to the PVR headquarters. Found 
subsequently to be prevalent in the area, with many heard, several seen 
and one trapped in both 1995 and 1996: a nest containing a single egg 
was discovered in a forest island in Ausust 1995. 


STRIOLATED PUFFBIRD Nystalus striolatus*” 


An individual noted at PSC on 15 August 1992 (White ez al. 1993) is 
apparently the second record for Beni, the first coming from humid 
Andean foothill forest (Serrania Pil6n) (Parker 1989). 


R.C. Brace & }. Hornbuckle 41 Bull. B.O.C. 1998 118(1) 


PALE-BREASTED SPINETAIL Synallaxis albescens 


This spinetail which has been found in low tangled scrub in both 
seasonally inundated and dry savanna (PVR and TRF estancias), 
particularly at the edges of forest islands and of the FFF, shows 
considerable, apparently undocumented plumage variation, which is 
assumed to be age-related. According to Ridgely & Tudor (1994), the 
tail should exhibit a dull brownish hue, but that of most birds examined 
(total of 51 trapped) displayed a rufous tinge, which can be marked and 
thus suggestive of Sooty-fronted Spinetail Synallaxis frontalis. 
Although some birds showed a pure rufous crown as expected, in the 
majority it was brown with some rufous speckling, and in a small 
number the crown was plain brown, variation which was apparent in 
skins scrutinized (BMNH). Note that the Plain-crowned Spinetail S. 
gujanensis, which has been trapped (three in 1995) in forest both at 
TRC and south of PVR, has more extensive rufous on the wings 
(remiges in addition to coverts), as is the case in S. frontalis and to some 
degree too in S. hypospodia, considered below. One _ individual 
controlled on 6 August 1994 was re-trapped in the same location (PVR 
forest island) on 23 August 1996. 


CINEREOUS-BREASTED SPINETAIL Synallaxis hypospodia* 


In comparison to S. albescens this species was less numerous, though 
observed routinely (up to c. five per day) in the peripheries of forest 
islands, at the edge of the FFF and in TRF scrub, requiring seemingly 
slightly taller (3+ m) vegetation. The rufous wing coverts were always 
duller in tone, and the black of the throat more extensive. It was found 
to be significantly heavier too: 14.9-18.1 (16.9) g, s.d. 1.04, n=17 as 
compared to 9.2—14.3 (11.2) g, s.d. 0.89, n=50 (Z= — 6.130, P<0.001). 
The two species are easily separated on song (see Ridgely & Tudor 
1994). One S. hypospodia caught (PRV scrub) on 6 August 1995 was 
re-trapped at the same site on 23 August 1996. (Re-trap weights have 
been excluded from the comparison presented above.) 


SPECKLED SPINETAIL Cranioleuca gutturata* 


An individual trapped at PSC on 12 August 1992 (White et al. 1993) 
was the first record for the BBS of this species, which has been found at 
a small number of other Benian localities (see Gyldenstolpe 1945, Davis 


et al. 1994). Additionally, several were seen and one mist-netted (17 
August) at TRC in 1994. 


PLAIN ANTVIREO Dysithamnus mentalis 


A specimen procured at the BBS in 1988 by Omar Rocha (see Davis 
et al. 1994) was the first from lowland Bolivia. However, it had been 
reported from Beni previously by Parker (1989), who found it to be 
fairly common above 800 m (Serrania Pil6n) and who heard the species 
also at low elevation, 20 km southeast of San Borja. We have observed 
it to be not uncommon at the BBS, with individuals often paired. A 
total of seven have been trapped: three in the FFF, three at 08 and one 
at TRC. 


R.C. Brace & $. Hornbuckle 42 Bull. B.O.C. 1998 118(1) 


STRIPE-CHESTED ANTWREN Myrmotherula longicauda**” 

One caught at FNC on 16 September 1992 (White et al. 1993) is the 
only record for Beni of this essentially foothill species (c. 500-1100 m) 
(Remsen & ‘Traylor 1989, Ridgely & ‘Tudor 1994). Prior to this 
sighting, the putative occurrence of Streaked Antwren M. surinamensis 
was deemed far more likely since this congener, which is very similar in 
appearance, occupies habitats more akin to those prevalent at the BBS 
than those frequented usually by M. longicauda, although admittedly 
M. surinamensis has only a limited and discontinuous distribution in 
northern and eastern Bolivia. 


SULPHUR-BELLIED TYRANT-MANAKIN WNeopelma sulphureiventer 

This relatively little known species occurs locally in southwest 
Amazonia (Brazil, Peru and Bolivia) (Gyldenstolpe 1945, Ridgely & 
Tudor 1994). Twelve individuals were secured at TRC (two being 
re-trapped in 1995), two were handled in the FFF and two mist-netted 
at 08. White ez al. (1993) trapped one bird at PSC and three at FNC. 
From experience of the species on the Rios Beni and Quizer in Beni and 
Santa Cruz, respectively, Remsen et al. (1988) describe it as being an 
inconspicuous resident of undergrowth of riverine forest, and 
commented that it is unclear whether or not the species is a bamboo 
specialist. Our work indicates that it is more catholic in its habitat 
preferences than the first of these two comments would suggest, and in 
the absence of any bamboo at the four ringing sites with which we are 
familiar, indicates that bamboo is not necessarily a habitat prerequisite. 


OCHRE-BELLIED FLYCATCHER Muonectes oleagineus** 

A total of eight Mionectes were mist-netted in seasonally inundated 
forest at TRC in 1995, over the 2-4 and 15-17 August periods, five 
being caught on one day. On the basis of the presence or absence of 
buffy/tawny tertial edgings (Ridgely & Tudor 1994, J. V. Remsen pers. 
comm.), four birds were identified firmly as M. oleagineus, and 
one—having no tertial edgings—as the sibling, McConnell’s Flycatcher 
M. macconnelli which had been recorded from the BBS (riverine forest) 
previously (Rocha 1988). Appraisal of the other three individuals 
remains inclusive, despite retrospective comparison with skins of the 
two species held by the BMNH. No Mionetes spp. were seen in either 
1994 or 1996, and none was observed either by White et al. (1993), who 
remained at Trapiche for over one week in 1992. M. oleagineus is 
widespread in lowland Bolivia (Arribas et al. 1995), and although it has 
been reported as favouring second growth and edge habitat and 
M. macconnelli described as inhabiting undisturbed forest (Willis et al. 
1978), the two species have been found also to occur together, in hilly 
upland forest and river-edge second growth (600m altitude) near 
Puerto Linares in La Paz department (Capparella & Lanyon 1985). 


SUIRIRI FLYCATCHER Swirivi suiriri* 

Three individuals of the northern form, S. s. affinis, treated 
separately sometimes as the Campo Suiriri (see Ridgely & Tudor 1994 
for discussion), have been recorded. One was seen in the periphery of a 


R.C. Brace & }. Hornbuckle 43 Bull. B.O.C. 1998 118(1) 
forest island in the TRF estancia on 30 August 1995 (RCB, JWP-H), 


and two were observed pursuing each other subsequently in fairly open 
savanna (PVR estancia) on 12 August 1996. Within Bolivia this 


subspecies is found in Beni and Santa Cruz (Remsen & Traylor 1983, 
Parker & Rocha 1991, Davis 1993). 


WHITE-CRESTED TYRANNULET Serophaga subcristata 


Over the 1994-96 period 12 trapped birds were identified as this 
species which is widespread, but identification of those (1=8) examined 
in 1994 is regarded now as being unreliable since it became apparent 
retrospectively in 1995 that the White-bellied Tyrannulet S. munda 
(see below)—a confusingly similar species—is found on the reserve 
also. Serpophaga spp. were seen on almost a daily basis on the PVR 
estancia in 1995 and 1996; the majority were assigned tentatively to 
S. subcristata. ‘Two individuals were re-trapped, one in 1995 (TRF 
estancia scrub) and one in 1996 (FFF), both in the same location as 
ringed. A specimen collected from forest island terrain 40 km east of 
San Borja on 30 August 1985 (Cabot 1990) was the first record of the 
species from Beni; Parker (1989) reported seeing three or four birds 
20—27 km east of San Borja (June) in thorny woodland and bushes. 


WHITE-BELLIED TYRANNULET Serpophaga munda* 


Of the total of nine Serpophaga secured (19-21 August 1995) in 
undergrowth at the edges of various forest islands or of the FFF, or in 
TRF scrub in 1995, five displayed a greyer, less olivaceous mantle and 
a paler belly (lacking or almost lacking any yellowish suffusion) than 
did the remainder, and were identified therefore as S. munda. That 
some S. munda can show traces of a yellow (possibly age-related) wash 
below necessitates that species separation must proceed cautiously (see 
Plain ‘Tyrannulet [nezia ornata below), though is aided greatly if their 
characteristic calls are heard (Parker 1989, Ridgely & Tudor 1994, pers. 
obs.). Apparently not recorded from Beni previously (Arribas et al. 
1995), though Parker (loc. cit.) indicated that he may have seen one 
27 km east of San Borja. The species occurs widely in Bolivia, but has 
yet to be recorded from Pando. Quite possibly S. munda visits the BBS 
solely during the austral winter, descending from Andean foothill and 
valley breeding areas (Ridgely & Tudor 1994). The putative migratory 
status of this species at the BBS, and of S. subcristata also, requires 
investigation. 


PLAIN TYRANNULET /nezia inornata 


This was easily the commonest tyrannulet encountered in the 
peripheries of forest islands, with a total of 25 being mist-netted on the 
PVR and TRF estancias. It was seen occasionally too in forest at TRC 
(with two birds trapped), where Serpophaga spp. were not encountered. 
The species is superficially very similar in appearance and behaviour to 
the two Serpophaga spp. dealt with above, and similarly great care is 
needed with field identification (see Ridgely & Tudor 1994). One 
individual ringed in 1995 was handled again in the same forest island in 


1996. 


R.C. Brace & }. Hornbuckle 44 Bull. B.O.C. 1998 118(1) 


WHITE-THROATED SPADEBILL P latyrinchus mystaceus 

Examination of birds trapped at TRC, in the FFF and in secondary 
forest 3 km south of the reserve (total of 20 individuals) showed that in 
common with three specimens secured by Rocha at the BBS previously 
(see Parker et al. 1991), they were somewhat ochraceous below, 
displaying intermediacy in coloration between that of lowland P. m. 
bifasciatus and of the Andean foothill form, P. m. partridgeiz. Such 
intermediate plumage substantiates the current subspecific status given 
to these forms whose ranges, together with that of P. m. zamorae from 
Peru and Ecuador, provide collectively an example of a circum- 
Amazonian distribution found in several bird groups (Remsen et al. 
1991). An individual collected just south of Trinidad (Schmitt & 
Schmitt 1987) was. bright yellow below, matching closely P. m. 
bifasciatus. ‘Two birds were re-trapped in 1995; one of these was 
handled for a third time in 1996, indicating residency within the FFF. 


CRESTED DORADITO Pseudocolopteryx sclateri** 

With the only other reports for this species in Bolivia relating to five 
males and two females acquired in December 1937 (Gyldenstolpe 
1945), detection of the species on the PVR estancia in August 1996 
was totally unexpected, though the site of those initial sightings, El 
Consuelo (Beni) to the east of Reyes, is only c. 100 km to the northwest 
of the BBS. It is pertinent to point out that the northern part of the 
distributional range of P. sclateri is characterized entirely by a 
scattering of highly restricted localities (Ridgely & ‘Tudor 1994). 
Twenty four individuals were trapped in the LGN cyperacean fringe, 
where the species was detected first by Joan and George Hardie on 
31 July (a party of four or five birds); singles were seen in two areas of 
savanna adjacent to curiches. Of those birds examined, nine were adult 
males, six were immature, and the remaining nine thought to be adult 
females. Six individuals were re-trapped (one twice), with four of the 
trap intervals being 14-17 days in length, thus demonstrating 
residency. Although confiding at times, the species was inconspicuous 
and it is possible therefore that it may have been overlooked previously. 


XENOPSARIS Xenopsaris albinucha* 

One obtained in TRF scrub on 22 August 1994 (JH) is the only 
record for the reserve. An additional sighting relates to an individual 
observed on 30 July 1995 at the edge of secondary forest 3 km south of 
the BBS (RCB, JWP-H); seen also at Trinidad (one on 22-23 July 
1996). The species appears to be scarce in Bolivia as elsewhere, though 
it has been found at a number of localities in Beni and Santa Cruz 
(Parker & Rowlett 1984) and has occurred also in Chuquisaca (Arribas 
et al. 1995). 


ASHY-HEADED GREENLET Hylophilus pectoralis* 

One trapped at TRC on 17 August 1995 is the first eicerel for Beni, 
although calls heard prior to this in winter 1995 near Riberalta 
(adjacent to the Brazilian border) by Sjoerd Mayer (pers. comm.), were 
probably of this species. It has a widespread Amazonian distribution 


R.C. Brace & }. Hornbuckle 45 Bull. B.O.C. 1998 118(1) 


(Sick 1993), and has been recorded in Pando (Gyldenstolpe 1945) and 
Santa Cruz (Noel Kempff Mercado National Park) (T. A. Parker & 
J. M. Bates unpubl. data). 


LONG-TAILED REED-FINCH Donacospiza albifrons* 


Within Bolivia, this species has been recorded only in Beni (Arribas 
et al. 1995). It was detected first in 1984, in open grassland in the 
vicinity of San Borja by Schmitt & Schmitt (1987), who obtained, 
additionally, two specimens 39 km west of Trinidad. It has been found 
26-30 km east of San Borja by Parker (1989), who observed family 
parties. New for the reserve in 1994, when a party of five was found in 
the PVR savanna (1 August); two were trapped subsequently in tall 
grass in the TRF estancia. Observations made in 1995 and 1996 have 
revealed that it occurs regularly in the area, with a further five sightings 
in both years (maximum of two birds seen). Moreover, five individuals 
were mist-netted (four in 1996). It is anticipated that in due course the 
species will be found in the department of Santa Cruz, since otherwise 
this western outpost implies a distributional jump of almost 1000 km 
from known localities in Brazil and Paraguay, spanning much 
seemingly suitable terrain, an unlikely circumstance. 


TAWNY-BELLIED SEEDEATER Sporophila hypoxantha* 


Known in Bolivia from Beni, Santa Cruz and La Paz (Arribas et al. 
1995), it was not recorded until 1995 when several males in breeding 
plumage were identified first by JWP-H (PVR estancia); two males 
were trapped subsequently in 1996. These trapped birds were with a 
flock of Sporophila spp., comprising many male Rusty-collared and 
Dark-throated Seedeaters S. collaris and S. ruficollis, and a small 
number of Double-collared and male Grey-and-Chestnut (Rufous- 
rumped) Seedeaters S. caerulescens and S. hypochroma, together with 
many unidentifiable birds. No doubt the species had been overlooked 
previously in such flocks, which are encountered not infrequently in the 
savanna during the July-September period. 


Acknowledgements 


We wish to acknowledge support from both Earthwatch and the Department of the 
Environment (U.K.) who sponsored our ‘Forest Islands of Bolivia’ expeditionary 
research from 1994 to 1996; funding from the latter was in the form of a Darwin 
Initiative grant. 

We thank especially Carmen Miranda of the Academia Nacional de Ciencias de 
Bolivia, who is Director of the BBS, for permission to undertake this work and for 
continuing help and encouragement. There are of course a plethora of other local people 
on whom we were dependent for logistical support and advice. They are too numerous to 
be referred to all individually, but the following should not escape specific mention: 
Sabina Stab and Roberto Urioste (former and current resident Scientific Coordinators at 
E] Porvenir), Alan and Erika Hesse of the Asociaci6n Armonia (Santa Cruz), Susan Davis 
and Tim Killeen of the Museo de Historia Natural ‘“Noel Kempff Mercado”’ (Santa 
Cruz), and Teresa and Celia Pérez (Trinidad). Contributors to field observations 
(including mist-netting) included Enzo Aliaga, Antonio Balderraguay, Mark Blazis, 
Karina Carrillos, Carolina Caceres, Betty Flores, Marcelo Hinojosa, Omar Martinez, 
Robin Mitchell, James Pearce-Higgins, Victoria Rojas, Dennise Quiroga, Gabriel 
Quisbert, André Rodriguez, Elva Villegas, and a number of the EarthCorps volunteers, 


R.C. Brace & }. Hornbuckle 46 Bull. B.O.C. 1998 118(1) 


in particular George and Joan Hardie who obtained the initial views of Accipiter 
superciliosus and Pseudocolopteryx sclateri at the BBS. Thanks go also to Sjoerd Mayer > 
(Cochabamba) for providing information on Hylophilus pectoralis. We acknowledge Jon 
Fleldsa (Centre for Tropical Biodiversity), and Carsten Rahbek and Jan Bolding 
Kristensen (Copenhagen Ringing Centre), all of the Zoological Museum of the 
University of Copenhagen, for providing the metal rings used in our studies. 

Finally, we thank Peter Colston of the Natural History Museum at Tring, where skins 
were examined (particularly of Synallaxis and Mionectes spp.). ; 


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Cambridge, U.K. 

Davis, S. E. 1993. Seasonal status, relative abundance and behavior of the birds of 
Concepcion, Departamento Santa Cruz, Bolivia. Fieldiana Zool. n.s. 71: 1-33. 
Davis, S. E., Rocha, O., Sarmiento, J. & Hannagarth, W. 1994. New departmental 

records and notes for some Bolivian birds. Bull. Brit. Orn. Cl. 114: 73-85. 

Flores, E. I. 1988. Perfil ornitologico de la reserva de la biosfera ““Estacion Boe 
Beni’. Mus. Natl. Hist. Nat. (Bolivia), Comunicacion No. 8: 7-14. 

Gyldenstolpe, N. 1945. A contribution to the ornithology of northern Bolivia. Kungl. 
Svensak Vetenskapsakad. Handl. Ser. 3, 23: 1-300. 

IUCN 1992. Protected Areas of the World: A Review of National Systems. Vol. 4 Nearctic 
and Neotropical. International Union for Conservation of Nature and Natural 
Resources. Gland, Switzerland, and Cambridge, U.K. 

Miranda, C., Ribera, M. O., Sarmiento, J., Salinas, E. & Navia, C. (eds) 1991. Plan de 
Manejo de la Reserva de la Biosfera, Estacion Biologica del Beni, 1991. Academia 
Nacional de Ciencias de Bolivia/Estacion Biolégica del Beni, La Paz. 

Parker, T. A. 1989. An Avifaunal Survey of the Chimane Ecosystem Program Area of 
Northern Bolivia, 17-26 June 1989. Unpublished report (for Conservation 
International). 

Parker, T. A. & Rocha, O. 1991. La avifauna del Cerro San Simon, una localidad de 
campo rupestre aislado en el depto. Beni, noeste Boliviano. Ecol. Bolivia 17: 15-29. 

Parker, T. A. & Rowlett, R. A. 1984. Some noteworthy records of birds from Bolivia. 
Bull. Brit. Orn. Cl. 104: 110-113. 

Parker, T. A., Castillo, A., Gell-Mann, G. & Rocha, O. 1991. Records of new and 
unusual birds from northern Bolivia. Bull. Brit. Orn. Cl. 111: 120-138. 

Remsen, ah V. & Traylor, M. A. 1983. Additions to the avifauna of Bolivia. Part 2. Condor 
85: 95-98. 

Bunter J. V. & Traylor, M. A. 1989. An Annotated List of the Birds of Bolivia. Buteo 
Books, Vermillion, South Dakota. 

Remsen, J. V., Schmitt, C. G. & Schmitt, D. C. 1988. Natural history notes on some 
poorly known Bolivian birds, Part 3. Gerfaut 78: 363-381. 

Remsen, J. V., Rocha, O., Schmitt, C. G. & Schmitt, D. C. 1991. Zoogeography and 
geographic variation of Platyrinchus mystaceus in Bolivia and Peru, and the 
circum-Amazonian distribution pattern. Orn. Neotrop. 2: 77-83. 

Ridgely, R. S. & Tudor, G. 1989. The Birds of South America: Vol.1. The Oscine 


Passerines. Univ. Texas Press. 


S.L. Olson 47 Bull. B.O.C. 1998 118(1) 
Ridgely, R. S. & Tudor, G. 1994. The Birds of South America. Vol. 2. The Suboscine 


Passerines. Univ. Texas Press. 

Rocha, O. 1988. Adicién de especies a la avifauna de la reserva de la biosfera “‘Estacion 
Bioldgica del Beni’, Bolivia. Ecol. Bolivia 12: 13-15. 

Rocha, O. 1990. Lista preliminar de aves de la reserva de la biosfera “‘Estacion Biologica 
del Beni’. Ecol. Bolivia 15: 57-68. 

Sarmiento, G. 1983. The savannas of tropical America. Pp. 245-288 in F. Bouliére (ed.), 
Tropical Savannas. Ecosystems of the World No. 13. Elsevier Scientific Publishing 
Company, Amsterdam, The Netherlands. 

Saunders, D. A. & Hobbs, R. J. (eds). 1991. The Role of Corridors. Chipping Norton, 
New South Wales, Australia. 

Schmitt, C. G. & Schmitt, D. C. 1987. Extensions of range of some Bolivian birds. Bull. 
Brit. Orn. Cl. 107: 129-134. 

Sick, H. 1993. Birds in Brazil: A Natural History. Princeton Univ. Press. 

Wege, D. C. & Long, A. J. 1995. Key Areas for Threatened Birds in the Neotropics. 
BirdLife Conservation Ser. No. 5. BirdLife International, Cambridge, U.K. 

Willis, E. O., Wechsler, D. & Oniki, Y. 1978. On the behavior and nesting of 
McConnell’s Flycatcher: does female rejection lead to male promiscuity? Auk 95: 
1-8. 

White, A. G., Brace, R. C., Duffield, G. E., Hesse, A., Payne, A. J. & Spick, S. 1993. 
Nottingham University Bolivia Project 1992: An Ornithological Survey of the Beni 
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Addresses: Robin Brace, School of Biological Sciences (Life Science), University of 
Nottingham, University Park, Nottingham NG72RD, U.K. Jon Hornbuckle, 5 
Grove Road, Sheffield S7 2GY, U.K. 


© British Ornithologists’ Club 1998 


Notes on the systematics of the Rockrunner 
Achaetops (Passiformes, ‘Timaliidae) and its 
presumed relatives 


by Storrs L. Olson 
Received 27 February 1997 


The Rockrunner or Damara Rockjumper Achaetops pycnopygius, which 
inhabits rocky country in Namibia and southwestern Angola, has had 
a rather curious taxonomic history. The species was “originally’’ 
described on three different occasions, first in the African sylviid genus 
Sphenoeacus as S. pycnopygius (Sclater in Strickland & Sclater 1852). It 
was next independently described as Drymoica (=Prinia) anchietae by 
Bocage (1868). Maintaining its association with the Sylviidae, Gray 
(1869) listed it as Megalurus pycnopygius. Finally, Sharpe (1869) 
unwittingly described the bird anew as Chaetops grayi. Hartlaub (1869: 
126) pointed out that this was the same species as Bocage’s Drymoica 
anchietae, and Tristram (1870: 497 footnote) showed that both of these 
were synonyms of Sclater’s name. He also quoted Jules Verreaux to the 
effect that the species ‘“‘cannot properly be included in any one of the 
genera to which it has been referred, and that probably a new genus 
should be established for its reception’’. This opinion notwithstanding, 


S. L. Olson 48 Bull. B.O.C. 1998 118(1) 


the bird was known at least until 1922 as Chaetops pycnopygius and was 
associated with the South African rockjumpers Chaetops frenatus and 
C. aurantius in the Timaliidae (e.g. Sharpe 1883). 

Roberts (1922: 227), a notorious generic splitter, paved the way for 
the eventual complete dissociation of pycnopygius from Chaetops by 
creating a new genus for it, Achaetops, on the basis of “‘its much shorter 
legs, and softer feathers on the crown’’. Nevertheless, Achaetops was 
still closely associated with Chaetops, and usually also with the Boulder 
Chat Pinarornis plumosus, in the family Timaliidae (e.g. W. L. Sclater 
1930), a treatment that continued through the first four editions of 
Roberts’ Birds of South Africa (Roberts 1940, McLachlan & Liversidge 
1957, 1970, 1978). 

Meanwhile, however, undercurrents arose that were to carry 
Achaetops and Chaetops off in different directions. These may be traced 
back to a few simple unsupported declarations by Delacour (1946: 11): 


Nous avons exclu du groupe des Timaliinés un certain nombre d’oiseaux africains qui 
y avaient été encore incorporés par W. L. Sclater [1930] et par D. A. Bannerman 
[1936]. Ce sont les espéces suivantes: Pinarornis plumosus est un Turdiné voisin de 
Cercotrichas podobe, apparenté sans doute aux Copsychus. Chaetops frenatus est un 
Traquet proche de Saxicola et de Cichladusa. Achaetops pycnopygius est un Sylviiné 
voisin des Melocichla . . . 


Not one of these associations has borne up under scrutiny (Olson 1984, 
1990, this study). It was thus Delacour who was responsible for 
Chaetops being placed in the Turdidae with the thrushes—I erred 
(Olson 1984) in crediting Ripley (1952) with being the first to do this, 
as he doubtless took his cue from Delacour. On the basis of its syrinx, 
Chaetops is definitely not a thrush (Olson 1984). Delacour’s suggestion 
of a relationship between Achaetops pycnopygius and the Moustached 
Warbler Melocichla mentalis is evidently what led White (1960: 20) to 
associate these two species with the Grassbird Sphenoeacus afer and 
to suggest “‘that their relationships would be better expressed by 
placing all three species concerned in the genus Sphenoeacus’’. Not long 
thereafter, what had once been three different genera became the 
““Sphenoeacus mentalis superspecies’’ (Hall & Moreau 1970: 159), a 
curious term considering that S. afer is the type species of the genus. 
This is an outstanding example of the evils of “‘compiler taxonomy’’, 
combined with abuse of the so-called superspecies concept, both of 
which have had a detrimental effect on modern ornithological 
systematics. Although White’s treatment was followed by numerous 
authors apart from Hall & Moreau, it is fortunate that recent influential 
works (e.g. Maclean 1985, Traylor 1986) have reverted to the use of 
three monotypic genera for these species. 

So we have seen the Rockrunner saltate from being congeneric with 
Chaetops, to a monotypic genus of Timaliidae, to a monotypic genus of 
Sylviidae, to congeneric with Sphenoeacus, to a superspecies with 
Melocichla mentalis and now back to being a monotypic genus of 
Sylviidae, with virtually no discussion of characters or the injection 
of new systematic information of any kind. Except for its generic and 
English names, the former association of this species with Chaetops has 
become totally obscured. 


S. L. Olson 49 Bull. B.O.C. 1998 118(1) 


Material examined 


Skeletons: Bradypterus luteoventris USNM 318312, USNM 318313; 
Dromaeocercus brunneus MRAC 50616; Amphilais (Dromaeocercus ) 
seebohmi USNM 432211; Melocichla mentalis UMMZ 208325, UMMZ 
218573; Achaetops pycnopygius "TM 32629; Chaetops frenatus USNM 
558653; Sphenoeacus afer USNM 558700, USNM 558701; Megalurus 
timoriensis USNM 561990, YPM 7089; Bowdleria p. punctata NMNZ 
22848: Pinarornis plumosus ROM 121100; Turdoides jardinett USNM 
558675. 


Results 


Examination of osteology of Sphenoeacus, Achaetops, and Melocichla 
discloses that these are sufficiently distinct from one another as to rule 
out any two of them as being congeneric. Sphenoeacus afer differs from 
the other two in the proportionately much shorter rostrum and 
premaxillary symphysis, the arched ridge of the dorsal nasal bar 
(culmen), narrower interorbital bridge, the distinctly notched and little 
inflated ectethmoid, and much broader and rounded zygomatic 
processes. The overall resemblance of the skull of S. afer is actually 
closer to the timaliid Turdoides than to either of the “‘sylviids’’ with 
which it has been allied. The manubrium of the sternum is much 
shorter in S. afer than in either Melocitchla or Achaetops. Although the 
skulls of S. afer and Melocichla are about the same size, the leg 
elements of S. afer are much smaller, and the distal wing elements are 
markedly more reduced, the carpometacarpus being about half the 
length of the ulna versus well over half in Melocichla. Compared to 
S. afer, the tarsometatarsus of Melocichla is longer and not as robust, 
and in Achaetops the tarsometatarsus and tibiotarsus are much longer 
and more slender, with the distal end not strongly curved and the 
plantar crest less ossified. The skull and mandible of Achaetops differ 
strikingly from Sphenoeacus or Melocichla in the very long, narrow bill, 
longer and more slender mandibular symphysis, and narrower frontal 
area. In these respects and in the morphology of the tarsometatarsus, 
Achaetops was identical to Chaetops. In fact, I could find no osteological 
differences apart from size by which these two “‘genera’’ could be 
distinguished. 

In plumage, Achaetops shares a light superciliary stripe and light 
malar stripe with Chaetops and also with Sphenoeacus and Melocichla. 
All but Chaetops have a black malar stripe as well, but this would be 
obscured in males of Chaetops, in which the entire throat is black. The 
breast streaks of Achaetops are seen in females of Chaetops (absent in 
Melocichla and only faintly indicated in Sphenoeacus). In both 
Achaetops and Chaetops the crown and back are heavily streaked (absent 
in Melocichia, back streaked but crown only faintly so in Sphenoeacus). 
Achaetops and Chaetops share a dark rufous belly that is absent in the 
other two genera, the rufous extending up onto the breast in Chaetops. 
They also share a strongly rufescent rump patch of loose, decomposed 
feathers, absent in the other genera. The pale tips to the rectrices of 
Achaetops (also in Melocichla) have become large white patches in 


S. L. Olson 50 Bull. B.O.C. 1998 118) 


Chaetops, which is also unique in having white tips to the secondary 
ccoverts. Interestingly, the remicle in both Chaetops and Achaetops has a 
white tip, lacking in the other two genera. 

In summary, the plumage of Chaetops differs from Achaetops in 
being strongly sexually dichromatic, in the more extensively rufous 
underparts, expansion of the white tips of the rectrices and the addition 
of white to the secondary coverts, and in the black throat of males. 
Although there is no real difference in the “softness”? of the crown 
feathers, the tarsometatarsus is proportionately longer in Chaetops (44% 
vs. 37% of wing length), as Roberts (1922) maintained, although such 
variation in tarsal proportions occurs commonly within numerous other 
accepted genera of birds. 

Both Chaetops and Achaetops are obligate inhabitants of rocky 
outcrops and are apparently quite similar in behaviour (Maclean 1985). 
It should be noted, however, that the Boulder Chat Pinarornis 
plumosus, another rock-dwelling passerine in southern Africa, is quite 
dissimilar in syrinx and osteology and appears to belong among the 
‘“‘proto-thrushes’’ including Myadestes, Neocossyphus, Stizorhina, and 
Modulatrix (Olson 1990). Because there were no grounds for 
dissociating Achaetops from Chaetops in the first place, and because a 
close relationship between Achaetops and either Sphenoeacus or 
Melocichla is not supported by osteology, there is no reason not to 
regard the similarities in plumage, osteology, and habits of the 
rockjumpers as indicative of relationship, with Chaetops being a larger, 
more ornately plumaged derivative of Achaetops. ‘This relationship is 
probably best expressed at the generic level, with Achaetops Roberts, 
1922, becoming a junior subjective synonym of Chaetops Swainson, 
1832. 

This brings us back to the question of the familial relationships of 
the re-expanded genus Chaetops. When I showed that the syrinx of 
C. frenatus was not thrush-like (Olson 1984), I merely suggested that 
the genus be returned to the Timaliidae, where it had nearly always 
been placed previously. On the other hand, ornithologists have been 
content for some time to accept C. pycnopygius as a warbler, so place- 
ment of the genus in the Sylviidae would seem equally plausible. 
Unfortunately, these are the two most ill-defined and problematical of 
the larger taxa of Old World passerines and no diagnostic characters 
have been identified that would permit a definitive decision to be made at 
this point. 

Irwin (1985: 99) concurred that Chaetops (sensu stricto) belonged in 
the Timaliidae, citing as diagnostic of that family a tail that is 
‘““moderately to well graduated with the outermost pair of rectrices 
sharply truncated and falling considerably short of the others”. This is 
not a convincing character, however, considering that numerous species 
of presumed Sylviidae have similar tails (e.g. Melocichla mentalis). As 
remarked by Irwin (1985), however, there are relatively few timaliids in 
Africa, and in southern Africa there is only the enigmatic Lioptilorms 
(Lioptilus auct.) and Turdoides, the latter being an Asian genus that has 
radiated secondarily in Africa. As he notes; Chaetops has no 
resemblance to either of these genera, as is also borne out by osteology. 


S.L. Olson 51 Bull. B.O.C. 1998 118(1) 


By contrast, the Sylviidae have radiated rather extensively in Africa, 
but likewise none of the African members of that family seem obviously 
related to Chaetops. 

Chaetops is very distinct osteologically from Turdoides, but that 
genus differs considerably from various other Timaliidae as well. As 
shown here, Chaetops is also very different osteologically from either of 
the genera of Sylviidae (Sphenoeacus and Melocichla) with which 
Achaetops has been associated. An informed solution to this systematic 
problem cannot be had without a great deal more study using various 
lines of evidence. Although some core group of Asian genera in the 
Timaliidae are probably monophyletic, the family has long been 
regarded as something of a wastebasket, so that the placement of an 
outlying genus in the Timaliidae carries with it a certain implicit 
ambiguity. For this reason, it is preferable to maintain Chaetops 
(including Achaetops) in the Timaliidae, rather than transferring it to 
the Sylviidae, which might convey a misleading impression of 
knowledge that we do not yet possess. 


Acknowledgements 


I am most grateful to the curators of the following institutions for lending specimens 
supplemental to those in the National Museum of Natural History, Smithsonian 
Institution, Washington (USNM): Transvaal Museum, Pretoria (TM); University of 
Michigan Museum of Zoology, Ann Arbor (UMMZ); Royal Ontario Museum, Toronto 
(ROM); National Museum of New Zealand, Wellington (NMNZ); Museum Royal de 
l Afrique Centrale, Tervuren (MRAC). 


References: 

Bannerman, D. A. 1936. Birds of Tropical West Africa. Vol. 4. Crown Agents for the 
Colonies, London. 

Bocage, J. V. B. du. 1868. Aves das possessOes portuguesas d’Africa occidental que 
existem no Museu de Lisboa. Jornal de Ciencias Mathematicas Physicas e Naturaes 
[Lisboa] 2: 38-50. 

Delacour, J. 1946. Les timaliinés. Ozseau 16: 7-36. 

Gray, G. R. 1869. Hand-List of Genera and Species of Birds, Distinguishing Those 
Contained in the British Museum. Part I. British Museum, London. 

Hall, B. P. & Moreau, R. E. 1970. An Atlas of Speciation in African Passerine Birds. 
British Museum (Natural History), London. 

Hartlaub, G. 1869. Bericht tiber die Leistungen in der Naturgeschichte der Vogel 
wahrend des Jahres 1868. Archiv fiir Naturgeschichte 35 (band 2): 105-132. 

Irwin, M. P. S. 1985. Chaetops and the Afrotropical Timaliidae (Babblers) Honeyguide 3: 
99-100. 

Maclean, G. L. 1985. Roberts’ Birds of South Africa. 5th edn. Trustees of the John 
Voelcker Bird Book Fund, Cape Town. 

McLachlan, G. R. & Liversidge, R. 1957. Roberts’ Birds of South Africa. Revised edn. 
Trustees of the South African Bird Book Fund, Cape Town. 

McLachlan, G. R. & Liversidge, R. 1970. Roberts’ Birds of South Africa. 3rd edn. 
Trustees of the John Voelcker Bird Book Fund, Cape Town. 

McLachlan, G. R. & Liversidge, R. 1978. Roberts’ Birds of South Africa. 4th edn. 
Trustees of the John Voelcker Bird Book Fund, Cape Town. 

Olson, S. L. 1984. Syringeal morphology and relationships of Chaetops (Timaliidae) and 
certain South African Muscicapidae. Ostrich 55: 30-32. 

Olson, S. L. 1990. Preliminary systematic notes on some Old World passerines. Riv. Ital. 
Orn. 59: 183-195. 

Ripley, S. D. 1952. The thrushes. Postilla 13: 1-48. 

Roberts, A. 1922. Review of the nomenclature of South African birds. Annals of the 
Transvaal Museum 8: 187-272. 


In Brief 52 Bull. B.O.C. 1998 118(1) 


Roberts, A. 1940. Birds of South Africa. H. F. & G. Witherby, London. 

Sclater, W. L. 1930. Systema Avium Afthiopicarum. Taylor and Francis, London. 

Sharpe, R. B. 1869. On the genus Chaetops. Proc. Zool. Soc. London 1869: 163-164. 

Sharpe, R. B. 1883. Catalogue of the Birds in the British Museum. Vol. 7. British Museum 
(Natural History), London. 

Strickland, H. E. & Sclater, P. L. 1852. List of a collection of birds procured by Mr 
C. T. Andersson in the Damara country in South Western Africa with notes. 
Pp. 141-160 in W. Jardine’s Contributions to Ornithology, part 7. 

Swainson, W. 1831 (=1832). Birds. Im Swainson and Richardson, Fauna Boreali 
Americana. Part 2. Longman et al., London. 

Traylor, M. A., Jr. 1986. [African Sylviidae] Jn E. Mayr & G. W. Cotrell (eds), Check-list 
of Birds of the World. Vol. 11. Museum of Comparative Zoology, Cambridge, 
Massachusetts. 

Tristram, H. B. 1870. Notes on some Old-World species of passerine birds. Ibis (2) 6: 
493-497. 

White, C. M. N. 1960. Notes on some African warblers. Bull. Brit. Orn. Cl. 80: 18-21. 


Address: Department of Vertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 


© British Ornithologists’ Club 1998 


IN BRIEF 


Differences in tarsal length between adult 
female Montagu’s and Pallid Harriers: 
an easy method to separate specimens 


by William S. Clark & Roger Clarke 


Received 2 April 1997 


Adult females of Pallid Harrier Circus macrourus and Montagu’s 
Harrier C. pygargus are often misidentified in museum collections as 
the other because of their similar plumages. Both have dark brown 
upperparts, buffy to creamy, heavily stréaked underparts, and similar 
tail patterns. Adult males and juveniles differ between species and are 
rather easy to distinguish. 

Some differences between the species, especially adult females, have 
been pointed out by Svensson (1971) and illustrated in Bruun et al. 
(1986). These differences, as well as some new ones, are summarized by 
Forsman (1995). However, none of these references mention the 
difference in leg lengths. 

One of us (WSC) noticed, from observing Pallid and Montagu’s 
Harriers perched on bare ground near each other, that Montagu’s 
appear to have much shorter legs, resulting in a more horizontal perch 
attitude, compared to the more upright stance of Pallid Harriers. 

To test if there was a clear separation between species of this measure 
that could be used as a species indicator, we measured the tarsal length 
of a large sample of adult female specimens in the British Museum 


In Brief 53 Bull. B.O.C. 1998 118(1) 
TABLE 1 


Tarsal length (mm) of adult 
female Montagu’s and Pallid Harriers 


BMNH AMNH 
C. pygargus 57.4-67.1 (26) 55.2-65.5 (16) 
(mean) 61.6 61.0 
C. macrourus 70.6—77.5 (27) 71.5—77.8 (18) 
(mean) 74.2 75.0 


(Natural History) (BMNH) and the American Museum of Natural 
History (AMNH). We used the standard measurement of tarsus as 
described in Baldwin et al. (1931). 

We found no overlap in the measure of tarsi between adult females of 
Montagu’s and Pallid Harriers (Table 1). 

The ranges of tarsi measurements given by Nieboer (1973) are 
55-65 mm (61) for pygargus and 63-76 mm (72) for macrourus. 'These 
are in general agreement with our measurements, but the overlap of 
ranges in his measurements and the lower mean macrourus suggest that 
his lower range of macrourus was due to one or more misidentified 
pygargus specimens. 

Even allowing for as much as a 5% measuring error, tarsal 
measurement clearly facilitates the correct identification of adult female 
specimens. 

Described differences ‘between the two adult females are useful in 
field identification but are less useful for identification of museum 
specimens. However, they can be used to check the identity based on 
tarsal measurement. These are: 

1. ‘The more distinct whitish facial ring of the Pallid Harrier, which 
extends across the throat of Pallid but not Montagu’s. 

2. Differences in markings on the secondaries; illustrated by Jonsson 
(1993). 

3. Differences in markings on axillaries and underwing coverts; 
pointed out for the first time by Forsman (1995). 


Although the difference in the position of the emargination on primary 
number 9, as illustrated on page 84 of Bruun ez al. (1986), also serves to 
distinguish the species, apparently it is not being used, as we have 
found many misidentified specimens. This method should be used to 
verify the identification made using the tarsal measurements. 

We found that tarsal measurements of. museum specimens are 
somewhat difficult, particularly locating the proper upper end of the 
tarsi with the caliper or divider. We took extra care in finding the 
proper measure points. 

Because raptors capture prey with their talons, it seems reasonable to 
hypothesise that tarsus length in each species has evolved with prey 
preferences. Apparent adaptations to detect prey by listening (Rice 


In Brief a4 Bull. B.O.C. 1998 118(1) 


1982) imply specialisation of the genus Circus to prey hidden amongst 
rank vegetation. Long tarsi would assist in reaching such prey, and so it 
has been suggested that differences in tarsus length proportionate to 
body size in the harriers are linked to the height of vegetation in their 
preferred hunting habitats (Nieboer 1977). For example, the Marsh 
Harrier Circus aeruginosus has long tarsi and forages in tall marsh 
vegetation. However, no clear distinction between Montagu’s and 
Pallid Harriers in habitat preference has yet been identified, but a 
marked contrast has been discovered in their prey preferences. Pallid 
Harriers most often hunt for passerines, whereas Montagu’s Harriers 
specialise more in lizards, large Orthoptera, and probably nest contents 
(Clarke 1996). 

In Accipiters, long tarsi occur in species that hunt for flying prey and 
shorter tarsi in those living on ground-dwelling prey (Wattel 1973). 
Long tarsi could give Pallid Harriers an edge in striking at fleeing birds. 
Adaptation to more agile prey is also evidenced by the greater reversed 
sexual dimorphism (Newton 1977) in this species as compared to 
Montagu’s Harriers, as well as their streamlining in wing shape and 
larger foot size. 

In summary, tarsal measurements are a quick and easy method to 


check the identity of all museum specimens of adult female Montagu’s 
and Pallid Harriers. 


Acknowledgements 


We thank the curators and collection managers of the American Museum of Natural 
History, especially Allison Andors, and the Natural History Museum (Tring) for 
assistance and permission to look at specimens of both harriers. R. Banks and J. Schmitt 
provided critical comments on earlier drafts. 


References: 

Baldwin, S. P., Oberholser, H. G. & Worley, L. G. 1931. Measurements of Birds. Sez. 
Publ. Cleveland Mus. Nat. Hist. Vol. 2. 

Bruun, B., Delin, H. & Svensson, L. 1986. Birds of Britain and Europe. Country Life 
Books, Twickenham. 

Clarke, R. 1996. Montagu’s Harriers. Arlequin, Chelmsford. 

Forsman, D. 1995. Field identification of female and juvenile Montagu’s and Pallid 
Harriers. Dutch Birding 17: 41-54. 

Jonsson, L. 1993. Birds of Europe, with North Africa and the Middle East. Princeton 
Univ. Press. 

Newton, I. 1979. Population Ecology of Raptors. T. & A. D. Poyser, London. 

Nieboer, E. 1973. Geographic and ecological differentiation in the genus Circus. Ph.D. 
dissertation. Free-University, Amsterdam, Netherlands. 

Rice, W. R. 1982. Acoustical location of prey by the Marsh Hawk: adaptation to 
concealed prey. Auk 99: 403-413. 

Svensson, L. 1971. Stapphok Circus macrourus och angshok C. pygargus—problemet att 
skilja dem at. Var Fagelvdrld 30: 106-121. 

Wattel, J. 1973. Geographical differentiation in the Genus Accipiter. Publ. Nuttall Orn. 
Club no. 13. 


Addresses: W. S. Clark, 7800 Dassett Court, Annandale, VA 22003, U.S.A. R. Clarke, 
New Hythe House, Reach, Cambridge CB5 0JW, U.K. 


© British Ornithologists’ Club 1998 


In Brief 55 Bull. B.O.C. 1998 118(1) 


Franklin’s Gull Larus pipixcan at South 
Georgia 


by Keith Reid 
Received 24 April 1997 


On 20 January 1997 at 18.45 (local time) an unusual gull was seen flying 
around Freshwater Bay, Bird Island (54°01’S, 38°03’W). When first 
noticed the bird was flying over the beach, after which it landed briefly 
on the shore before flying off to the southeast and out of sight. The initial 
impression was of a small buoyant gull, compared to the resident Kelp 
Gull L. dominicanus, recalling Black-headed Gull L. ridibundus, with 
noticeable long dark wings and a distinct dark marking on the head. 

It was watched for approximately 5 minutes in good light through 
7 X 42 binoculars at a range down to 20 m and the following description 
taken. Head—forehead, lores, chin and throat white; crown, nape 
and ear-coverts dark, forming half hood extending to just in front 
of eye. Eye dark with distinct pale upper and lower eye crescents. 
Bill dark and approximately equal in length to the loral distance. 
Upperparts—mantle and wing coverts dark grey, outer primaries 
darker and lacking pale mirrors. Inner primaries and secondaries with 
pale tips forming whitish trailing edge contrasting with dark secondary 
bar and outer primaries. ‘l'ail white with distinct black sub-terminal 
band. Underparts—white. 

In examining the possible species involved it is apparent that this 
bird belonged to one of the group of small gulls which have dark 
head markings in adult plumage. Within this group three species 
which regularly occur in South America—Brown-hooded Gull Larus 
maculipennis, Andean Gull L. serranus and Grey-headed Gull L. 
cirrocephalus—can all be ruled out as they have much paler grey mantle 
and wing-coverts which form a very characteristic wing pattern 
(Harrison 1983), different from the bird in question. This leaves 
Laughing Gull L. atricilla and Franklin’s Gull L. pipixcan, both of 
which have a much darker grey base colour across the wings and mantle. 
Although the Laughing Gull shows a distinct dark subterminal tail 
band in first winter plumage it has generally grey, not white, under- 
parts (Grant 1982). In addition the Laughing Gull’s bill is longer than 
the loral length, often appearing “‘heavy and drooping”’ (Grant 1982). 
Franklin’s Gull has a distinctive dark half hood with white eye-crescents 
in all plumages and first winter birds show a distinct dark subterminal 
tail band (Grant 1982). From this combination of plumage and struc- 
tural characters the bird was identified as a first winter Franklin’s Gull. 

Other than Kelp Gull, which is a common resident, Dolphin Gull L. 
scoresbu is the only other gull species confirmed on the South Georgia 
list (Prince & Croxall 1996). A single record of Brown-hooded Gull was 
considered unacceptable by Bourne (1988) as the description published 
(Delany et al. 1988) could apply to a second winter Franklin’s Gull. 
Although Brown-hooded Gulls breed in southern South America, 


In Brief 56 Bull. B.O.C. 1998 118(1) 


including the Falkland Islands, they disperse north after breeding 
(Harrison 1983). Franklin’s Gull, however, nests mainly on inshore 
marshes in the prairies of North America and migrates south to winter 
along the Pacific coast of South America as far south as Valdivia, Chile 
(Harrison 1983). The highly migratory nature of this species is 
illustrated by previous records in the South Atlantic, Indian Ocean and 
Australia (Higgins & Davies 1996). In the South Atlantic there are 
single records from Tristan da Cunha in February 1956 (Swales & 
Murphy 1965) and the Falkland Islands, the South Orkney Islands and 
in the Scotia Sea (summarised in Prince & Croxall 1996). Watson 
(1975) refers to a single bird reported from Gough Island in February 
1956, citing Swales & Murphy (1965), and records from both Gough 
Island and Tristan da Cunha are reported in the review of records by 
Higgins & Davies (1996). Clearly both references relate to the single 
bird on Tristan da Cunha, and there do not appear to be any authentic 
published records from Gough Island. 


I thank J. P. Croxall, P. A. Prince and W. R. P. Bourne for their comments and advice 
and Carl Zeiss UK for the loan of binoculars. 


References: 

Bourne, W. R. P. 1988. Was it a Brown-hooded or a Franklin’s Gull at South Georgia? 
Sea Swallow 37: 64. 

Delany, S. N., Edwards, D. V. & Williams, T. D. 1988. Brown-hooded Gull Larus 
maculipennis: first record for South Georgia. Bull. Brit. Antarct. Surv. 78: 53-54. 

Grant, P. J. 1982. Gulls: A guide to identification. T. A. A-D. Poyser, Calton. 

Harrison, P. 1983. Seabirds: An identification guide. Christopher Helm, London. 

Higgins, P. J. & Davies, S. J. J. F. (eds) 1996. Handbook of Australian, New Zealand and 
Antarctic birds. Vol. 3: Snipe to Pigeons. Oxford Univ. Press. 

Swales, M. K. & Murphy, R. C. 1965. A specimen of Larus pipixcan from Tristan Da 
Cunha. [bis 107: 394. 

Prince, P. A. & Croxall, J. P. 1996. The birds of South Georgia. Bull. Brit. Orn. Cl. 116: 
81-104. 

Watson, G. E. 1975. Birds of the Antarctic and sub-Antarctic. American Geophysical 
Union, Washington, D.C. 

Woods, R. W. 1988. Guide of the Birds of the Falkland Islands. Anthony Nelson, 
Oswestry. 


Address: British Antarctic Survey, High Cross, Madingley Road, Cambridge CB3 OET, 
U.K. 


The date of publication of Montin’s 
description of Lagopus mutus 


by Tommy Tyrberg 
Received 26 April 1997 


The original description of the Ptarmigan Lagopus mutus was published 
by Lars Montin in Vol. 1 of Physiographiska Sdlskapets Handlingar 


In Brief 57 Bull. B.O.C. 1998 118(1) 


(Proceedings of the Physiographic Society). This volume (the only 
one to appear) was actually published in four sections (comprising 
pp. 1-64, 65-132, 133-220 and 221-318 respectively) over a period of 
ten years (Dal 1996), but only the first section bears a date (1776), 
which has until now been considered the date of publication of Lagopus 
mutus. 

Montin’s paper comprises pp. 150-155 and was therefore part of the 
third section, something which is obvious from the paper itself since 
the pages are marked “‘Del I. St. 3” (Part I Section 3).-The date of 
publication of this section can be determined from the minutes of the 
Physiographic Society. At the meeting on 2 May 1781 the Secretary 
announced that “‘nu 3:dje stycket af Handlingarne pa Herr Assessor 
Gjoérwells forlag war fardigt tryckt’’ (‘“‘the printing of the third section 
of the Proceedings at Mr Gjérwell’s Press is now finished’’), while at 
the next meeting on 3 October: “‘Secreteraren upwiste tredje stycket af 
Salskapets handlingar, hwilket war fran Stockholm nedkommit, sedan 
sista sammankomst”’ (““The Secretary displayed the third section of the 
Society’s Proceedings which had arrived from Stockholm since the 
previous meeting’’) (cited after Gertz (1940, pp. 15-16)). The date of 
publication is confirmed by a review in Ubpfostrings-Sdlskapets 
Tidningar (no. 32 p. 125 (26 April 1781)). That the Proceedings could 
be reviewed in Stockholm on 26 April, but had not yet arrived in Lund 
(where the Physiographic Society was situated) on 2 May is not 
surprising considering the distance (c. 600 km) and the slowness of 
communications during the eighteenth century. 

From these data it is clear that the date of publication for Lagopus 
mutus is actually 1781, and that the name should therefore be cited as 
Lagopus mutus (Montin, 1781). 


References: 

Dal, B. 1996. Sveriges zoologiska litteratur. En beradttande édversikt om svenska zoologer och 
deras tryckta verk 1483-1920. Orbis Pictus, Kjuge. 

Gertz, O. 1940. Kungl. Fysiografiska Sdallskapet 1 Lund 1772-1940. Historisk éverblick och 
Personfoérteckningar. Hakan Ohlssons Boktryckeri, Lund. 

Montin, L. 1781. Tvanne Arter af Snéripan. Physiographiska Salskapets Handlingar Del 
1.St. 3: 450-155. 


Address: Kimstadsvagen 37, S-610 20 Kimstad, Sweden. 


Wattled Cranes in Guinea-Bissau 


by N. F. Collar 


Received 30 April 1997 


Remarks by Hazevoet (Bull. Brit. Orn. Cl. 117: 56-59) concerning the 
handling in Collar & Stuart (1985, Threatened Birds of Africa) of a 
record of Wattled Crane Bugeranus carunculatus from Guinea-Bissau 
strike me as over-critical. As far as Collar & Stuart (1985) are 


In Brief 58 ~ Bull. B.O.C. 1998 118(1) 


concerned, this record did not: require his reinstatement: if we are 
guilty of “‘less than adequate’’ interpretation, so is he, for his assertion 
that we “cast doubt on the validity of the record’’ is unfounded. We 
wrote: ‘“‘A specimen, considered immature, was collected in 1948 at 
Lake Cufada, Fulacunda, in Guinea-Bissau’’, and went on exactly as 
quoted in Hazevoet’s first paragraph, concluding by citing Snow’s 
(1978) judgement that “‘it seems certain that the birds involved were 
either escapes or vagrants’ 

In no way, then, was this record doubted, and I cannot see why 
Hazevoet includes either Snow (1978) or Collar & Stuart (1985) amongst 
the unconvinced, or why he lumps Snow (1978) and Dowsett (1993) 
with Collar & Stuart (1985) in accusing them of failing to understand 
Frade & Bacelar’s Portuguese, which 1s a remark only really dirigible at 
me (I drafted the account in Collar & Stuart), and one which I reject. 
What is true is that my comment concerning “‘an incomplete reference 
that suggests the species had previously been recorded at this locality” 
was made without checking the bibliography in Frade & Bacelar (1955). 
In accumulating many references on the Wattled Crane for the account 
in Threatened Birds of Africa | had, under considerable time pressure, 
merely taken a copy of the pages dealing with the crane in their paper, 
and therefore did not refer to any of its sources. 

On re-reading these pages, however, I see that one particularly bald 
statement.in Frade & Bacelar, also quoted by Hazevoet (“J. A. Ferreira, 
in his article on the ‘Fauna of the Cufada Reserve’, mentions the species 
under the designation ‘grows de caruncula ’’), is clearly the source of my 
mistaken remark, as it can easily be misinterpreted (without mistrans- 
lation) as (a) suggesting that this work 1s not in the bibliography (other- 
wise the authors might just have written “‘Ferreira [1948], in his ...’’), 
hence my view of its citation as incomplete; and (b) implying that the 
species might well have been found in the reserve on several occasions 
(otherwise it is not clear why Frade & Bacelar, having already given the 
record Hazevoet is discussing, should have troubled to make this extra 
citation at all). Moreover, since only seven lines earlier they refer to 
‘““Major Araujo Ferreira’ as supplying the specimen evidence, and on the 
next page credit him (albeit there as Captain Araujo Ferreira) for the 
photographs, it is not even obvious that their author “J. A. Ferreira” (a 
common enough name in Portuguese, almost Smith in English!) is the 
same person, which compounds the illusion of multiple records. 

All this emphasises the importance of remaining alert to potential 
alternative readings of the literature, and of the value of tracing every 
reference for original rather than second-hand evidence. However, even 
Hazevoet may concede that the pursuit of a source as obscure as Bol. 
Cult. Guiné Port. (1948) in order to seek further details of evidence 
already accepted as valid may not represent optimal use of deadlined 
time in documenting a species with 110 other references to process, and 
with 176 other species jostling for treatment. 


Address: BirdLife International, Wellbrook Court, Girton Road, Cambridge CB3 O0NA, 
U.K. 


In Brief 59 Bull. B.O.C. 1998 118(1) 


Neotype of the Hooded Plover Charadrius 
rubricollis Gmelin, 1789 


by Ian A. W. McAllan & Leslie Christidis 
Received 17 Fune 1997 


Examination of the Australian bird literature reveals the continued use 
of two different species-group names for the Hooded Plover: 
Charadrius rubricollis Gmelin, 1789 and Charadrius cucullatus Vieillot, 
1818. Since the publication of the 1975 edition of the Australian 
Checklist (Condon 1975), most references in the Australian literature 
have adopted rubricollis (note that Condon also used the name in 1968). 
In the previous 75 years the name cucullatus was widely used in the 
Australian literature. Condon based his decision on what he stated was 
Oberholser’s “‘rejection’’ of Vieillot’s name in 1919. This is, however, 
not strictly true. Oberholser did not reject cucullatus; he resurrected 
rubricollis instead. 

Oberholser’s resurrection of rubricollis derives from Mathews’ 
discussion on the background of the name. Mathews (1913) showed 
that Gmelin’s description of rubricollis was simply a Latin translation of 
a description of a bird (in English) by Latham in volume III of his 
General Synopsis of Birds. This bird, called by Latham the ‘‘Red- 
necked Plover’’ was “‘Found in Adventure Bay, Van Diemen’s Land”’ 
and was evidently based on drawings of birds delineated on Captain 
Cook’s third voyage by the artist William Ellis (drawings now held in 
The Natural History Museum, London). One of these, number 67, is 
clearly that of a Hooded Plover (see for example Whitley 1970, wherein 
there is a monochrome photograph of this drawing). Latham’s 
description also recorded that the Red-necked Plover had ‘‘on each side 
of the neck a large square chestnut spot, the size of a silver penny,’ and 
“‘a little mixture of white around the bastard wing’’, characters not 
found in the Hooded Plover, nor in Ellis’ drawing 67. Mathews 
determined that Latham’s bird was also based in part on Ellis’ drawing 
number 63, a picture of a Red-necked Phalarope Phalaropus lobatus 
(Linnaeus, 1758), which had the relevant features. On the basis of 
Latham’s description and the evidence of the drawings, Mathews 
considered that the name rubricollis was not valid because it was based 
on two different species. 

Oberholser’s decision to resurrect the name involves the incorrect 
argument that Latham’s description was based primarily on the 
Hooded Plover. However as the name rubicollis refers to more than one 
taxon it would appear to be a case of instant homonymy and is thus not 
valid. 

Today the species-group name rubicollis is applied almost universally 
to the Hooded Plover. It has been used in many widely read 
publications such as Peters (1934), Condon (1975), Marchant et al. 
(1986), Sibley & Monroe (1990), Marchant & Higgins (1993) Collar 
et al. (1994) and Christidis & Boles (1994). However, the name 


In Brief 60 Bull. B.O.C. 1998 118(1) 


cucullatus has been used several times since the publication of Condon’s 
Checklist in 1975, principally in publications by the late G. M. Storr 
(in particular Storr, 1986a, b, 1987, 1991, Storr & Johnstone 1979, 
1988). In these works there is no discussion on the reason for using 
cucullatus. 

Given that there is still potential confusion with these names, and 
that the issue of the incorrect description has never been resolved, we 
think that it would be most appropriate for stability and universality of 
usage that the type of rubricollis be defined more precisely. The original 
descriptions were based on two drawings. Under the I.C.Z.N. (1985) 
the type specimens of a description based on illustrations are the 
animals described and not the illustrations themselves (Art. 72c (v)). 
The original specimens can no longer be located. Some specimens were 
known to have been associated with the corresponding drawings of 
Ellis, but a specimen of a Hooded Plover was not amongst these 
(Stresemann 1950). In any case there are very few bird specimens in 
existence of any species that can be proved to have been collected on 
any of Cook’s voyages (Whitehead 1969). 

Thus in this situation no lectotype can be made. Although we are 
reluctant to assign a neotype on the chance, albeit unlikely, that the 
original type can be located, it is essential for this taxon that its name be 
stabilised. The Hooded Plover is considered a threatened species 
(Collar et al., 1994) and its name has been used in legislation of the 
Australian States in which it is found. We therefore put forward 
the specimen HLW 5741 located in the H. L. White collection in the 
Museum of Victoria as the neotype. It is an adult male collected near 
Burnie on the north coast of ‘Tasmania on 13 January 1906 by W. H. 
Moffitt. This specimen was selected as it was collected near the original 
type locality. Ellis’ bird was from Adventure Bay on Bruny Island, 
within a few kilometres of mainland ‘Tasmania. The type locality now 
becomes the locality at which the neotype was collected (vide I.C.Z.N. 
1985 Art. 75f), that is, near Burnie on the north coast of ‘Tasmania. 


Acknowledgements 


The reasons for neotypification were reviewed by Dr R. Schodde of the Standing 
Committee on Ornithological Nomenclature of the International Ornithological Congress 
and W. E. Boles of the Taxonomic Advisory Committee of the Royal Australasian 
Ornithologists Union. 


References: 

Christidis, L. & Boles, W. E. 1994. The Taxonomy and Species of Birds of Australia and 
its Territories. Royal Australasian Ornithologists Union Monograph 2. 

Collar, N. J., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to Watch 2. The world list of 
threatened birds. BirdLife International, Cambridge (U.K.). 

Condon, H. T. 1968. A Handlist of the Birds of South Australia. 2nd edn. South 
Australian Ornithological Association, Adelaide. 

Condon, H. T. 1975. Checklist of the Birds of Australia. Part 1, Non-passerines. Royal 
Australasian Ornithologists Union, Melbourne. 

Gmelin, J. F. 1789. Systema Naturae. 13th edn. G. E. Beer, Liepzig. 

I.C.Z.N. (International Commission on Zoological Nomenclature). 1985. International 
Code of Zoological Nomenclature. 3rd edn. International Trust for Zoological 
Nomenclature, London. 


In Brief 61 Bull. B.O.C. 1998 118(1) 


Latham, J. 1785. A General Synopsis of Birds. Vol. III(1). Leigh & Sotheby, London. 

Marchant, J., Prater, P. & Hayman, T. 1986. Shorebirds, an identification guide to the 
waders of the world. Christopher Helm, Bromley, Kent. 

Marchant, S. & Higgins, P. J. (eds). 1993. Handbook of Australian, New Zealand and 
Antarctic Birds, Vol. 2. Oxford Univ. Press. 

Mathews, G. M. 1913. The Birds of Australia. Vol. 3, Part 2. Witherby & Co., London. 

Oberholser, H. C. 1919. The status of Charadrius rubricollis Gmelin. Auk 36: 279. 

Peters, J. L. 1934. Check-list of Birds of the World. Vol. 2. Museum of Comparative 
Zoology, Harvard. 

Sibley, C. G. & Monroe, B. L. Jr. 1990. Distribution and Taxonomy of the Birds of the 
World. Yale Univ. Press. 

Storr, G. M. 1986a. Birds of the Houtman Abrolhos, Western Australia. Rec. W. Aust. 
Mus. Suppl. No. 24. 

Storr, G. M. 1986b. Birds of the south-eastern interior of Western Australia. Rec. W. 
Aust. Mus. Suppl. No. 26. 

Storr, G. M. 1987. Birds of the Eucla Division of Western Australia. Rec. W. Aust Mus. 
Suppl. No. 27. 

Storr, G. M. 1991. Birds of the South-west Division of Western Australia. Rec. W. Aust. 
Mus. Suppl. No. 35. 

Storr, G. M. & Johnstone, R. E. 1979. Field Guide to the Birds of Western Australia. 
Western Australian Museum, Perth. 

Storr, G. M. & Johnstone, R. E. 1988. Birds of the Swan Coastal Plain and adjacent seas 
and islands. Rec. W. Aust. Mus. Suppl. No. 28. 

Stresemann, E. 1950. Birds collected during Capt. James Cook’s last expedition 
(1776-1780). Auk 67: 66-88. 

Whitehead, P. J. P. 1969. Zoological specimens from Captain Cook’s Voyages. 7. Soc. 
Bibiogr. Nat. Hist. 5: 161-201. 

Whitley, G. P. 1970. Early History of Australian Zoology. Royal Zoological Society of 
N.S.W., Sydney. 


Addresses: 1. A. W. McAllan, 46 Yeramba St., Turramurra, New South Wales 2074, 
Australia. L. Christidis, Museum of Victoria, 71 Victoria Cres., Abbotsford, Victoria 
3067, Australia. 


© British Ornithologists’ Club 1998 


Books Received 62 Bull. B.O.C. 1998 118(1) 
BOOKS RECEIVED 


Snow, D. W. & Perrins, C. M. (eds) 1998. Birds of the western Palearctic, Concise 
Edition. Pp. 1694, 594 plates in colour, maps and text figures. Oxford University Press. 
ISBN 0-19-854099-X. £150. 26 x 22 cm. 2 Vols. in slip case. 


When I was an enthusiastic youngster (the adjective still applies!), the principal 
ornithology text, Witherby’s Handbook of British birds, was beyond the reach of my 
pocket. The publication in 1952 of The popular handbook, followed in 1962 by its 
companion on rarer birds, thankfully allowed me access to a condensed, but nevertheless 
invaluable, account of the identity and lives of the birds around me. 

A similar evolutionary pathway has now been followed by the successor to Witherby’s 
Handbook, Oxford University Press’s Handbook to the birds of Europe, the Middle East, and 
North Africa, generally known as the Birds of the western Palearctic, or BWP. The 9 volumes 
of BWP were published over a 17-year period. This period encompassed a massive prolifer- 
ation in the ornithological scientific literature, major changes in the distribution and numbers 
of some birds, a huge increase in the number of bird-watchers, and a greatly increased 
propensity of these to travel to see birds. Thus by the time volume 9 appeared in 1994, the 
species accounts in volume 1 were beginning to look distinctly aged. However, the time taken 
to produce the 9 volumes was an inevitable consequence of BWP’s scope, giving the latest 
comprehensive information on the lives of the birds of the region in some 7000 pages. 

For many, the detail of the information in BWP has proved too much, and its cost will 
render it beyond the reach of many who were unable to collect the volumes as they 
appeared over the years. A need was therefore felt for a shorter version, leading to the 
production of this Concise Edition, in two volumes and a mere 1700 pages! But these 
volumes are far more than a condensed version of BWP. 

The shortening of species accounts has entailed a complete re-write; during this 
process data have been up-dated, incorporating new taxonomic considerations and 
up-to-date information on distribution, movements and populations. This ciearly 
required an extensive revision of the maps, all of which have been re-drawn and 
presented in colours which allow easier interpretation than in BWP. 

In bringing the Concise Edition up-to-date over 80 species, not included in BWP, had to 
be added, necessitating new or revised artwork. Of the 594 colour plates, 231 are new or 
revised, and they are dispersed in the text so that most accompany the species accounts; 
however, this has not always been possible (e.g. the Varied Thrust Zoothera naevia is 
illustrated with the Northern Mockingbird Mimus polyglottos and Eastern Phoebe Sayornis 
phoebe, about 100 pages away from the Varied Thrush account and, more important, from 
plates of its relatives). The overall quality of the plates is excellent and, although the 
postures of some birds are suspect, they are designed to show off important distinguishing 
features. If only pipits would display themselves in the field as depicted on p. 1091, and 
waders as on p. 623! I found an error on p. 790 involving the transposition of the labelling 
of the Sooty and Bridled Terns Sterna fuscata and S. anaethetus. 

What has been lost from BWP? Sections headed Plumages, Bare parts, Moult, 
Measurements, Weights and Structure have been omitted (although wing length and 
weight ranges are given), as have the plates of eggs. All other sections are of course much 
reduced, especially those detailing Food, Social pattern and behaviour, and Voice (there 
are no sonograms). Nevertheless, Field characters, when used in conjunction with the 
improved plates, are adequate for identification and especially distinction from similar 
species. Sections on Distribution and Population are up-to-date and incorporate much 
recent information from eastern Europe. A major sacrifice throughout the book, but 
especially important here where changes in range, status and population size are 
described, has been the omission of references to original sources of data. 

Although those who need in-depth information on species biology will still need access 
to BWP, those who want to identify west Palearctic birds from descriptions and colour 
illustrations, to know where they occur, when and where they breed, and their status and 
population trends, will find their needs amply met in the Concise Edition. The volumes are 
attractively produced, but more important is the content, which is clearly presented and 
concise, and information should be readily located thanks to indexes of vernacular names 
in eight European languages. David Snow and Chris Perrins are to be congratulated on the 
production of this fine work, and these editors are clearly concerned at the precedent set by 
BWP, in that no-one calls it by its full title; they suggest that the Concise Edition should be 
called BWPC! I am concerned that young ornithologists may be prevented from owning 


Books Received 63 Bull. B.O.C. 1998 118(1) 


copies by the cost, but with the launch of these volumes in January 1988, there is plenty of 
time for them to request parents to start saving now for next Christmas! 


Chris Feare . 


Mikhailov, K. E. 1997. Avian eggshells: an atlas of scanning electron micrographs. 
Pp. 88, 16 monochrome plates, 36 text figures. British Ornithologists’ Club Occasional 
Publication No. 3. ISBN 0-9522886-2-1. £35. 30.5 x 21.5 cm. 


This book is not for the faint-hearted! Part of the difficulty in assessing work of this 
kind is that it is such uncharted territory; Dr Mikhailov’s work is so specialised that it is 
difficult, even for those with considerable knowledge of eggs, to comment meaningfully 
on the correctness of his results. The work is in two parts: the first discusses avian 
eggshell structure in general terms, making comparisons with the structure found in 
other vertebrates; the second, which constitutes the main part of the “‘atlas’’, consists of 

‘detailed descriptions of the vertical structure of eggshells in each of the major avian 
orders, suborders and families which Mikhailov has been able to examine. These are 
accompanied by excellent line diagrams, and microscopic photographs of cross sections of 
individual representative eggs. The photographs are beautifully reproduced, but to the 
uninitiated,they are likely to be difficult to interpret; I certainly found them to be so. As 
a result, the general reader will probably have to take the conclusions on trust. This is not 
to decry Dr Mikhailov’s achievement, as his work is highly important. 

While the results of this work are often at variance with the conclusions of Sibley and 
Ahlquist, they do not depart in any material way from the “traditional”? arrangement of 
Peters’ Check List, and lend support to a number of taxonomic relationships that were 
long suspected. Mikhailov shows, for example, that the emus and cassowaries are more 
closely related to each other than to other ratites, and likewise that the ostriches and 
elephant birds, with their branched pores, are probably close to each other and not to 
other ratites. The suborder Ciconii, as set up by Sibley and Ahlquist, is not supported by 
Mikhailov’s results, but support is given to the long-held but unconfirmed view that the 
Shoebill is an aberrant Pelecaniform. On the other hand, eggshell data do support a close 
relationship between the cathartids and the storks, and the Accipitridae and Pandionidae 
appear close to each other but not to other diurnal birds of prey. 

Michael Walters 


Mearns, Barbara & Richard 1998. The Bird Collectors. Pp. xvit472, numerous 
text-figures. ISBN 0-12-487440-1. Academic Press. £29.95. 25 x 17 cm. 


This book—beautifully produced, well illustrated and readable—has one of its aims to 
present a reasoned defence of what the authors describe as a now “‘generally vilified”’ class 
of ornithologists, those who built up the collections now residing in the world’s museums. 
_ It is also much more than that. The bird collectors whose lives and ornithological careers 
are described are a varied and fascinating lot; they are here brought to life (a few are still 
alive) by detailed accounts of their exploits, with a wealth of apt quotations from their 
diaries, journals and letters, and many old photographs and other illustrations. 

The 12 middle chapters devoted to the collectors, which make up the bulk of the book, 
classify them under headings such as ““Army Officers’’, ““Clergymen and Missionaries’’, 
“The Professional Field Collectors’, “The Great Accumulators” (Lord Rothschild is an 
easy first in this category), thus imposing some order though it is admitted that many 
collectors could be treated under more than one heading. The decision whom to include or 
exclude cannot always have been easy; some clearly had to be included, others were more 
borderline. In the main the list seems well chosen, except for the omission of any mention of 
Admiral Lynes, surely one of the outstanding collectors, unique in his day for collecting with 
a particular object in mind, to work out the taxonomy of a very difficult and species-rich 
genus (Cisticola), and notable too for his detailed, meticulous labelling of specimens. 

Three preliminary chapters include a survey of the historical and cultural background, 
followed by a useful discussion of methods of collecting and preserving specimens, and the 
place of collecting in the development of scientific ornithology. A concluding chapter 
discusses the importance of collections today, stressing their continued use by ornithologists 
and the varied data obtainable, and now often only obtainable, from museum specimens. A 
12-page Appendix lists the world’s largest collections of study skins, with numbers of 
specimens held, main geographical areas covered, and other points of interest. 

This is a book not only for the museum worker but for all who are interested in the 
history of ornithology. 

D. W. Snow 


Books Received 64 Bull. B.O.C. 1998 118(1) 


Juniper, T. & Parr, M. 1998. Parrots: a guide to the parrots of the world. Pp. 584, 88 
colour plates. Pica Press. ISBN 1-873403-40-2. £35. 24.5 x 18 cm. 


This latest guide from Pica Press follows earlier models in its general format, but deals 
with a large family of attractive birds of great commercial importance and partly as a 
result of this, of considerable conservation concern. The book recognises 352 species of 
parrot, and their classification is based on that of Sibley & Monroe (1990, 1993), with 
minor amendments. The species, and many gender, age and racial variants, are illustrated 
in the 88 plates by five artists. Introductory sections outline evolutionary relationships, 
classification, natural history, conservation status, threats and captive breeding. 
The species accounts provide information under the headings of identification, voice, 
distribution and status, ecology, description, sex/age, measurements, geographical 
variation and references, and a distribution map. In view of the international trade in 
parrots, the authors state that the book is designed to meet the identification needs of 
both field ornithologists and those involved in monitoring the movements and legal status 
of birds in the captive bird trade; for some species the identification section gives 
characters that may be used to identify birds in the field and in the hand. 

The introductory sections are disappointing and largely without references; in fact of 
the 9 references, 2 are not included in the reference list and one is incorrectly cited. The 
section on natural history is especially poor and here it is difficult to know what are 
established facts and what are the authors’ views. Examples of unsupported statements 
include: pre-roosting calling and intense interaction ‘may be linked to... foraging 
effectiveness’; ‘some species “‘appoint’”’ (authors’ quotes) sentinels to watch over feeding 
flocks’; ‘pairs that know each other well and have experience of one another breed more 
successfully’; there “may be a linkage between the onset of laying and incubation 
behaviour and the daytime darkness experienced by birds spending time in their 
nest-hollows’. The sections on conservation and threats are more substantial (but still 
largely un-referenced) and make important point, e.g. emphasising the political, social 
and economic basis of forest loss, a major threat to many parrot species. The authors 
recognise the socio-economic need for exploitation of parrots in some places, but also 
stress the need for improvements in welfare during transportation; this consideration 
could also have been extended to the conditions of parrots in their final homes as caged 
birds. 

The species accounts are concise and accompanied by clear distribution maps; these are 
monochrome, but as most parrots are non-migrants colour is unnecessary. Measure- 
ments, given as ranges, are largely taken from Forshaw (1989) but sample sizes are not 
given, so that without reference to this publication it is not possible to assess the 
reliability of these data. In the account of one of the species with which I am familiar, the 
Seychelles race of the Black Parrot Coracopsis nigra, the authors recognise the existence of 
the race (barkly1) and describe it as critically endangered, but include a surprising and 
unsupported statement that the species was possibly introduced to the Seychelles! One of 
the references at the end of this species account is not cited in the reference list. 

With plates by five artists, variation in style is to be expected. Most are typical of many 
post-Peterson field guides and are of high quality and attractive; those by Dan Powell are 
especially pleasing as, in addition to displaying identification features, his birds appear 
wonderfully alive. A helpful feature is the appearance of a drawing of a budgerigar on the 
caption page facing each plate, to show the relative sizes of the birds depicted. 

The index lists vernacular names, specific and subspecific names, but SAP aCe does 
not provide an alphabetical listing of generic names. 

Overall, this book is an attractive addition to the series and well undoubtedly be useful 
for those who need to identify parrots, though somewhat heavy to be taken into the field 
(along with all the other family guides!). For those who need more detailed information, 
however, this will have to be sought elsewhere and the references provided by this book 
may be of limited help. 

Chris Feare 


NOTICE TO CONTRIBUTORS 

Papers are invited from Club Members or non-members, especially on 
taxonomic and distributional topics; descriptions of new species are especially 
welcome and may be accompanied by colour photographs. Two copies of 
manuscripts, typed on one side of the paper, double spaced and with wide 
margins, should be sent to the Editor, Prof. Chris Feare, 2 North View 
Cottages, Grayswood Common, Haslemere, Surrey GU27 2DN, UK. All 
contributions, including In Brief articles, should follow the style of main 
papers in this issue of the Bulletin. 


A contributor is entitled to 10 free offprints (16 if 2 or more authors) of the pages of the 
Bulletin in which his contribution, if one page or more in length, appears. Additional 
offprints or offprints of contributions of less than one page may be ordered when the 
manuscript is submitted and will be charged for. Authors may be charged for proof 
corrections for which they are responsible. 


MEMBERSHIP 

Only Members of the British Ornithologists’ Union are eligible to join the 
Club, and to receive (postage free) four quarterly issues of the Bulletin, and the 
annual index, for an annual subscription of £12 (or U.S. $26). Applications, 
enclosing the annual subscription, should be made to the Hon. Secretary 
(address as below). 

The 1998 List of Members, and addresses will not be published with the 
Bulletin this year, but copies are available, on application (with a remittance 
of £1.00 to cover costs of production and postage), to the Hon. Secretary. 
Please advise the Hon. Secretary, without delay, of any address changes, or 
corrections, for despatch of the Bulletin. 


NON-MEMBER SUBSCRIBERS & APPLICATIONS FOR BACK 
NUMBERS OR OTHER PUBLICATIONS 

The Bulletin (for 1998 onwards), together with annual index, may be 
purchased (postage free) by Non-member Subscibers on payment of an annual 
subscription of £22 (or U.S. $45) on application to The Publications Officer, 
S. J. Farnsworth, Hammerkop, Frogmill, Hurley, Maidenhead, Berks SL6 5NL, 
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PAYMENTS 

All amounts quoted are net and should be paid in £ sterling, if possible. 
Payments in other currencies must include a further £4 for UK bank charges 
(except for annual rates in U.S. dollars, which are inclusive). All cheques or 
drafts should be made payable to the British Ornithologists’ Club. If preferred, 
remittances may be made by bank transfer direct to the Club’s bank 
account—Barclays Prime Account, Dale House, Wavertree Boulevard, 
Liverpool L7 9PQ, U.K. (Sort Code 20-00-87 Account No. 10211540), with 
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Bickley, Bromley, Kent BR1 2LF. 


CORRESPONDENCE 
Correspondence on membership, changes of address and all other matters 
should be addressed to the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA, U.K. For details 
of Club Meetings see inside front cover. 


Registered Charity No. 279583 


CONTENTS 


CLUB NOTICES 2.) 060/006 Sis aoccd Sia ed ein oe, aoe als ae ae 
JESsuP, L. ‘“Gibsone’s nondescript’ and the lost type of Scissirostrum dubium 
(Latham) o.oo eS a el, Ia et ota 
FRITH, C. B. & FRITH D. W. Hybridization between Macgregor’s Bowerbird 
Amblyornis macgregoriae and the Streaked Bowerbird A. subalaris 
(Ptilonorhynchidae) of New Guinea... .. 0... .... 9 J) eee 
MAURICIO, G. N. & DIAS, R. A. Range extensions and new records for forest 
birds in southern Rio Grande do Sul; Brazil... .... ... ee 
STILES, F. G. Notes on the biology of two threatened species of Bangsia 
tanagers in northwestern Colombia... ...+. 2.0.0.5: ©. eee 
FIGUEROA-ESQUIVEL, E. M., NAVARRO, S., A. G. & POZO-DE LA TIJERA, C. New 
distributional information on the birds of southern Quintana Roo, 
MEXICO: oe BES he ae aa ale te oe 
BRACE, R. C. & HORNBUCKLE, J. Distributional records of and identification 
notes on birds of the Beni Biological Station, Beni, Bolivia ......... 
OLSON, Ss. L. Notes on the systematics of the Rockrunner Achaetops 
(Passeriformes, Timaliidae) and its presumed relatives............. 

In Brief CLARK, W. S. & CLARK, R. Differences in tarsal length between adult 
female Montagu’s and Pallid Harriers: an easy method to 

separate specimens ...... 25... +++ s/s 5 5 

REID, K. Franklin’s Gull Larus pipixcan at South Georgia. ..... 

TYRBERG, T. ‘The date of publication of Montin’s description of 

Lagopus mutus.. 0 0. 6 ee cee io ss oe 

COLLAR, N. J. Wattled Cranes in Guinea-Bissau ............. 

McALLAN, I. A. W. & CHRISTIDIS, L. Neotype of the Hooded Plover 
Charadrius rubricolis Gmelin, 1789 ......-.... 925 eee 

BOOKS) RECEIVED ¢ 20s ss gala, sc 6 6 6% «obey qlee «keane « ote 


59 
62 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 


mail, after their current subscription has been paid. 


COMMITTEE 
Revd T. W. Gladwin (Chairman) (1997) Miss H. Baker (1994) 
Mrs A. M. Moore (Vice-Chairman) (1997) Dr R. P. Pr¥s-Jones (1995) 
Cdr M. B. Casement, OBE, RN N. H. F. Stone (1995) 
(Hon. Secretary) (1996) D. Griffin (1997) 


D. J. Montier (Hon. Treasurer) (1997) N. J. Redman (1997) 


Hon. Editor: Prof C. J. Feare 
Chairman of Publications Sub-committee: Mrs A. M. Moore 
Publications Officer: S. J. Farnsworth 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


3 ISSN 0007-1595 
pD 


Bulletin of the 


British Ornithologists’ Club 


Edited by 
Prof CHRIS FEARE 


Volume 118 No. 2 June 1998 


FORTHCOMING MEETINGS 


Tuesday 15 September 1998. Dr Nigel Collar, of BirdLife International, will speak on 
“Two Indian Enigmas’’, based on work he has done jointly with Dr Pamela C. Rasmussen, of 
the Smithsonian Institution, on the Forest Owlet Athene blewitti, (recently rediscovered partly 
as a consequence of this work). Nigel began his ornithological career with the Great Bustard 
Trust, then served as editor of the Birds of the Western Palearctic, before moving to his current 
position as Red Data Book compiler with BirdLife (formerly ICBP). 

Applications to the Hon. Secretary by 1 September, please. 


Tuesday 13 October 1998. Dr Ben Hatchwell will speak on “The cooperative 
behaviour of Long-tailed Tits”. After reading Zoology at Oxford (1981-84), he worked on 
population and biology and coloniality of Common Guillemots on Skomer Island, Wales, for 
his PhD at Sheffield, and subsequently moved to Cambridge to carry out research on the 
mating systems and parental behaviour of Dunnocks and Alpine Accentors. He became 
Lecturer in Zoology at Sheffield, in 1993, from where he has been studying Long-tailed Tits. 
His main interests in ornithology concern breeding systems, in particular the evolution of social 
behaviour and the conflicts of interest that exist between males and females. He has also 
conducted research on more applied topics, including the importance of woodlands for 
farmland birds, and the population biology of vulnerable seabirds. 

Applications to the Hon. Secretary by 29 September, please. 


Tuesday 17 November 1998. Dr W. G. (Bill) Porteous will speak on “Birds of the 
Humboldt Current’’. Bill was born and raised in Shetland, and may therefore be unique in 
having added Booted Warbler to his life list before he added Blue Tit. He subsequently 
qualified as a geologist, and this has given him the opportunity to pursue birds in various parts 
of the world, particularly in north and south America. He recently spent five years in Colombia, 
which provided some welcome exposure to the neotropics, which had long been a particular 
interest of his. The opportunity to participate in a voyage southwards along the coasts of Peru 
and Chile in late 1995 provided the material we are to see this evening. 

Applications to the Hon. Secretary by 3 November, please. 


Meetings are held in the Sherfield Building of Imperial College, South Kensington, London 
SW7. The nearest Tube station is at South Kensington, and car parking facilities are available; a 
map of the area will be sent to members, on request. The cash bar is open from 6.15 p.m., and 
a buffet supper, of two courses followed by coffee, is served at 7.00 p.m. (A vegetarian menu 
can be arranged if ordered at the time of booking.) Informal talks are given on completion, at 
about 8.00 p.m. 


Overseas Members visiting Britain are particularly welcome at meetings. For 
details in advance, please contact the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1998 


Apart from single copies made for the purposes of research or private study, or criticism or 
review, as permitted under UK law, no part of this publication may be reproduced, stored 
or transmitted in any form or by any means, except with prior permission in writing of the 
publishers, or in accordance with the terms of licences issued by the Copyright Licensing 
Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; for 
address see inside back cover. 


65 Bull. B.O.C. 1998 118(2) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 118 No. 2 Published 19 June 1998 


CLUB NOTES 


The eight hundred and seventy third meeting of the Club was held on Tuesday, 20 
January 1998 at 6.15 p.m. 36 Members and 11 Guests attended. 

Members present were: The Rev. T. W. GLapDWwIN (Chairman), 5S. P. de 
ALBUQUERQUE, Miss H. Baker, P. J. BeLMan, I. R. BisHop, D. M. Brap ey, D. R. 
Caper, Cdr. M. B. CASEMENT RN, Professor R. J. CHANDLER, Professor R. A. CHEKE, 
Professor C. J. FearE, G. D. FreLp, F. M. GAuNTLETT, A. Gipss, D. GRIFFIN, C. A. 
HELM, J. A. JoBLiINnG, R. H. Ketrve, M. B. Lancaster, Dr C. F. Mann, D. J. MONTIER, 
Mrs A. M. Moore, R. G. Morcan, Mrs M. N. MULLER, P. J. OLiver, M. L. PALinc, Dr 
R. P. Pryce-Jones, N. J. RepMan, J. E. RicHarpson, R. C. SELF, P. J. SELLAR, R. E. 
SHARLAND, Dr D. W. Snow, N. H. F. Stone, C. W. R. Storey and R. W. Woops. 

Guests attending were: Mrs G. BonHaM, Dr I. BuRRows (Speaker), Mrs J. B. CALDER, 
Mrs C. R. CasEMENT, Mrs J. M. GLapwin, Mrs M. H. GaAuntTLETT, Ms C. Horr, Mrs 
M. Montier, P. J. Moore, Mrs M. OLiver and Mrs A. Woops. 

On completion, Ian Burrows spoke on the Birds of Papua New Guinea, illustrated with 
an impressive selection of his own photographic slides. New Guinea is well known for its 
Birds of Paradise and Bowerbirds, and is recognised as a major centre for the adaptive 
radiation of many genera including the Fruit-pigeons Ptilinopus and Ducula, 
Owlet-nightjars Aegotheles and several kingfisher and honeyeater genera. This has 
resulted in almost 400 species endemic to the New Guinea region, with over 350 of these 
present in Papua New Guinea and its satellite islands. 

Ian’s talk focused on some of the major biogeographic zones of Papua New Guinea, 
with particular reference to the common birds present and, where relevant, some of the 
rare or unusual species endemic to each area. 

The volcanic island of Lihir in New Ireland Province provided the scene for a brief 
description of the breeding behaviour of the Melanesian Scrubfowl Megapodius eremita 
which uses geothermal energy to incubate its buried egg. The precocious fledgling digs its 
way to the surface after 30 days untended incubation and, shortly after reaching the 
surface, is fully independent and able to fly short distances. 

Tench Island, again in New Ireland Province, illustrated the large seabird colonies 
which may be found on remote atolls. Many thousands of White-capped Noddies Anous 
minutus, Brown Noddies A. stolidus and Sooty Terns Sterna fuscata breed here with 
minimal human interference. 

The avifauna within a 50 km radius of Port Moresby is probably the most extensively 
studied in Papua New Guinea. Major habitats include savannah, mangroves, freshwater 
marshes, lowland rainforest and forested foothills. Away from the forested areas, the 
avifauna is dominated by Australian species. Common breeders include Rainbow 
Lorikeet Trichoglossus haematodus, Rainbow Bee-eater Merops ornatus and Fawn- 
breasted Bowerbird Chlamydera cerviniventris, whilst Sacred Kingfisher Halcyon sancta 
and Shining Bronze-Cuckoo Chrysococcyx lucidus are migrants from the south. The 
freshwater marshes and lagoons support large numbers of herons and egrets Ardeidae 
and are vital refuelling sites for at least 23 species of transient Palearctic wader. 
Sharp-tailed Sandpiper Calidris acuminata, Pacific Golden Plover Pluvialis fulva and 
Red-necked Stint Calidris ruficollis are the commonest migrants. 

The lowland and foothill rainforest contains a diverse variety of endemic species and 
this was illustrated by some 20 slides of typically representative species including 
Common Paradise Kingfisher Tanysiptra galatea, Blue Jewel-babbler Ptillorrhoa 
caerulescens, Emperor Fairy-wren Malurus cyanocephalus, Golden Monarch Monarcha 
chrysomela, King Bird of Paradise Cicinnurus regius and Raggiana Bird of Paradise 
Paradisaea raggiana. 

The vast sago swamps of the Gulf Province have a similar range of species as the 
rainforests around Port Moresby, but their remoteness and low human population 


Books Received 66 Bull. B.O.C. 1998 118(2) 


density results in an abundance of the large ‘“‘game’’ birds such as Southern Cassowary 
Casuarius casuarius and Southern Crowned Pigeon Goura scheepmakeri. A method of 
luring these species, and the rarely seen New Guinea Flightless Rail Megacrex inepta, 
into 50 mm lens range was described. 

In contrast, the Bensbach river floodplain in Western Province resembles the 
Northern Territories of Australia, reflected by impressive numbers of resident Brolga 
Grus rubicunda, Australian Bustard Ardeotis australis and Black-necked Stork 
Ephippiorhynchus asiaticus, and up to 20 000 migrant Little Curlew Numenius minutus. 
The rare endemic Fly River Grassbird Megalurus albolimbatus also occurs there and the 
slides shown may be a photographic first. 

Stark contrast was provided by the Owen Stanley mountains in Central Province. 
Many species in the hill forest zone are endemic and are altudinal replacements for their 
lowland congeners. Slides of 10 representative species were shown, including 
Brown-headed Paradise Kingfisher Tanysiptera danae by its nest. Most species in the 
lower montane forest are endemic, and different from those of the lowlands. Typical 
species shown included Hook-billed Kingfisher Melidora macrorrhina, Hooded Pitohui 
Pitohui dichrous (the first bird shown to be poisonous), New Guinea Harpy-eagle 
Harpyopsis novaeguineae, Feline Owlet-nightjar Aegotheles insignis (Plate 1), Macgregor’s 
Bowerbird Amblyornis macgregoriae and Magnificent Bird of Paradise Cuicinnurus 
magnificus. ‘The high mountain forests and alpine grasslands of Mount Victoria host a 
small range of almost wholly endemic species. Typical examples shown included Papuan 
Lorikeet Charmosyna papou, Painted ‘iger-Parrot Psittacella picta, Mountain 
Owlet-nightjar Aegotheles albertsi, Greater Ground-Robin Amalocichla sclateriana, 
Crested Bird of Paradise Cnemophilus macgregori and Macgregor’s Bird of Paradise 
Macgregoria pulchra. 

The speaker concluded with a range of Papua New Guinea men in full tribal regalia 
showing plumes from many different species of Bird of Paradise and other endemic 
species. 


Erratum 

In the review by Michael Walters of Mikhailov’s book Avian 
eggshells: an atlas of scanning electron micrographs (Bull. B.O.C. 118(1): 
63), the final sentence should read ‘On the other hand, eggshell data 
do not support aclose relationship between the cathartids and the 
storks 


BOOK RECEIVED 


Barlow, C. and Wacher, T. 1997. A Field Guide to Birds of The Gambia and Senegal. 
Illustrated by A. Disley. Pp. 400. 48 colour plates, maps, text figures. Pica Press, ISBN 
1-873403-32-1. £26. 22 x 15 cm. 


This is a first class book which will be tremendously useful and popular with the large 
number of birders who visit the Gambia. It will also be very useful throughout West 
Africa as proven on a recent trip to Nigeria where it was greeted with great enthusiasm. 

The text is informative, giving the salient identification features with a most useful 
section explaining differentiation from the most similar species. The plates are 
well-organised covering 618 species and the illustrations for most of the species are good 
and capture the stance of the birds. Details of voice and habitat preference are 
comprehensive with much information not previously published. Status and distribution 
sections are up to date including the most recent information from Senegal. Taxonomic 
treatment is modern and the very comprehensive Indigobird section will be particularly 
appreciated. With such a well-produced book it seems churlish to mention a fault but the 
record of Adamawa Turtle Dove from Djoudj and attributed to me is an inexplicable 
error. 

This book is a credit to the authors and the publisher and should be bought by anyone 
travelling to West Africa and also to anyone who has an interest in the birds of the region. 

S. J. R. Rumsey 


Plate 1 


4. 


Plate 2 


1. Head of recently dead Amaurornis magnirostris Karakelong Island, September 1996. 
2. Recently dead A. magnirostris, Karakelong Island, September 1996. 

3a. Comparison of A. magnirostris type specimen with specimens of A. olivaceus and 
A. moluccanus underparts. 

3b. Upperparts. 

4. Comparison of bill structure and length of A. magnirostris and A. olivaceus. 
Photographs by Frank Lambert. 


F.R. Lambert 67 Bull. B.O.C. 1998 118(2) 


A new species of Amaurornis from the ‘Talaud 

Islands, Indonesia, and a review of taxonomy 

of bush hens occurring from the Philippines 
to Australasia 


by Frank R. Lambert 
Received 7 February 1998 


Bush hens in the Talaud Islands 


During August and September 1997, I observed and heard several 
forms of rail, whose identity was puzzling, on the island of Karakelong 
in the Talaud archipelago, Indonesia. One of these proved to be a new 
species of Gymnocrex (Lambert 1998), whilst a second is the subject of 
this paper. 

The first observation was made on 17 August, when a rallid was 
heard giving very quiet single alarm notes on a steep slope in primary 
forest, inland from Toabatu in the vicinity of Mt Manuk (4°19’N 
126°50’E). This observation was made at an altitude of 300-400 m a.s.1. 
in an area of understorey dominated by rattan Calamus sp., in 
undisturbed habitat. The bird was very inquisitive, and approached to 
within a few metres but was never seen well. However, it was clearly 
large and appeared to be a species of Amaurornis: I had already heard 
the distinctive grating and squealing calls typical of a bush hen, 
believed to be Plain Bush Hen A. olivaceus, near the coast on 15 August 
1996. 

The presence of a bush hen on the island was of interest in itself, 
since no previous records of A. olivaceus had been reported for the 
Talaud Islands by White and Bruce (1986), although F. G. Rozendaal 
(in litt. 1997) had heard bush hens on the island during a visit in 1985. 
The brief views of the bird obtained on 17 August did not suggest a 
typical bush hen, the bird appearing to be considerably larger and 
darker than individuals of A. olivaceus moluccanus that I had seen 
elsewhere in Indonesia (This taxon is treated as a separate species, 
A. moluccanus, by some recent authors, including Sibley & Monroe 
(1990)). Furthermore, the habitat was perhaps unusual, since in 
Indonesia A. olivaceus is usually encountered in secondary growth and 
adjacent damp grasslands rather than in primary forest (pers. obs.). In 
some parts of its range, however, it has been found in swampy forest 
bordering wetlands (Rand & Gilliard 1967) and in forest edge (Beehler 
et al. 1986). 

Three days later, another individual of this Amaurornis was 
observed, again in tall primary forest inland from Beo, but this time in 
a relatively flat area with scattered patches of very swampy ground and 
a relatively sparse understorey, at c. 100 m altitude. I disturbed the bird 
and it ran about 20m before stopping abruptly. Here it extended its 
neck upwards and stood motionless, evidently watching me. A clear 


FR. Lambert 68 Bull. B.O.Cx 199848) 


MINDANAO 

° MIANGAS 

100 200 300 ra SARANGANI ISLAND 
ISLAND 


Scale in Km. 


ta He 5 
KAWIO ISLANDS NENUSA 
; KARAKELONG ISLANDS 

TALAUD ISLANDS 
Beo, 


Rainis 


saueoaauy 


KADARUANE 
hehe 


Biitiitisii) 
50 25. «50 
Scale in Km. 


Figure 1. Location of the Talaud Islands. 


impression of the bird was obtained: it was very robust, all dark and 
had a striking long, broad, deep-based pale greenish bill and a red eye. 

On 2 September, an Amaurornis was observed three times, in forest 
edge near the coast, to the south of Beo (see locality map, Figure 1). 
Calls presumed to be bush hens had been heard in the area at first light, 
but later some other unusual calls were heard that at first were 
mistakenly believed to have been made by a frog. They had been 
coming from a wet scrubby area on the edge of forest where there was 
permanent water. The same calls were also heard originating in forest 
beside a road and, apparently in response, from a narrow strip of 
littoral swamp forest bordered by coconut plantation on the other side 
of the road. 

Using playback of these frog-like vocalisations, an Amaurornis 
resembling that observed on 20 August was again observed, albeit very 
briefly, as it approached quietly from the swamp forest in response to 
the tape. The bird was exceedingly shy and burst into a run before 
binoculars could even be raised. A slightly longer view was obtained of 
a different individual from a hidden vantage point behind a tree, again 
using playback. Lastly, on 2 September, a pair was briefly observed 
again, without using playback, as they moved along the edge of a small 
stream covered with scrub and patches of trees but bordering more 
open dense herbage and grassland. 

Despite the brevity of these accumulated views, it was clear that this 
was a different taxon to the Plain Bush Hens that the author had 
observed previously in the Moluccas, and on the island of Sangihe in 
the previous month. 


F. R. Lambert 69 Bull. B.O.C. 1998 118(2) 


Subsequently, on 6 September 1996, a man was encountered selling 
various rallids in Beo. These had been caught near the small town of 
Rainis, and included one dead specimen of the unidentified 
Amaurornis, three live and one dead Plain Bush Hen Amaurornis 
olivaceus moluccanus, as well as one live Gymnocrex rail (Lambert 1998). 
After discussion with officers from the local office of the Directorate 
of Forest Protection: and Nature Conservation (PHPA), the dead, 
unidentified Amaurornis was purchased from the hunter and a 
specimen was made. 

The presence of two species of Amaurornis bush hen on Karakelong 
is of great significance, and provides good reason to review the 
taxonomy of the bush hens that occur from the Philippines to 
Australasia. The White-breasted Waterhen Amaurornis phoenicurus, 
which also occurs in this region, 1s not considered a bush hen and is not 
dealt with in this review. 


Amaurornis taxonomy 


There have been two views of the taxonomy relating to the Amaurornis 
bush hens that occur in the Philippine archipelago, the Moluccan 
Islands, New Guinea region and Australia. Apart from the very 
distinctive Isabelline Bush Hen A. isabellinus of Sulawesi, the majority 
of recent authors, presumably following Peters (1934), have treated all 
the “‘plain” bush hens from Philippines to Australasia as one species: 
Plain Bush Hen A. olivaceus (e.g. Olsen 1973, Ripley 1977, White & 
Bruce 1986, Dickinson et al. 1991, Andrew 1992, Marchant & Higgins 
1993). These authors did not discuss any alternative treatment. 

A. olivaceus of the Philippines was originally described as a species 
(Meyen 1834) and was retained as such by Sharpe (1894) in his 
catalogue of the birds in the British Museum, vol. 23, which was the 
first attempt at a classification of the family. Sharpe also listed A. 
moluccanus, with a range covering the Moluccas, New Guinea, 
Bismarck Archipelago and Australia. In contrast to other authors, 
McAllan & Bruce (1988) also recognised two species, treating those in 
the Philippines as Plain Bush Hen A. olivaceus, but other populations, 
from the Moluccas to Australia, as a separate species, Rufous-tailed 
Bush Hen A. moluccanus. This latter taxonomic treatment was 
subsequently adopted without discussion by Sibley & Monroe (1990) 
and del Hoyo et al. (1996), despite the fact that no author had ever 
provided adequate reason for recognising two species. McAllan & 
Bruce (1988), for example, simply stated that “‘the populations of 
bush-hen in Australasia are quite distinct from A. olivaceus (Meyen 
1834) of south east Asia (White & Bruce 1986) and may best be treated 
as an allospecies, as adapted by us here (see also Wolters 1982)’’. 

In contrast, the recent authoritative and more taxonomically rigorous 
checklist to birds of the Indomalayan region (Inskipp et al. 1996) 
treated these taxa as conspecific (as Plain Bush Hen A. olivaceus), 
following Marchant & Higgins (1993) and specifically mindful of the 
fact that no publication had provided a reasoned account of why two 
separate allopatric species should be recognised (T. Inskipp, pers. 


F.R. Lambert 70 Bull. B.O.C. 1998 118(2) 


comm.). However, the existence of two similar, sympatric Amaurornis 
taxa on Talaud clearly provides a good reason to re-examine the 
taxonomy of these birds. 


How many species of bush hen? 


Bush hens from the Philippine Archipelago are consistently larger than 
those to the south-east (i.e. Moluccas to Australasia). Indeed, my 
measurements of the series of Amaurornis from the Philippines, 
Moluccas, New Guinea, New Britain and Australia in the Natural 
History Museum (BMNH), Tring (Table 1), and those published 
elsewhere (Table 2), show that olivaceus from the Philippines is 
longer-billed, longer-winged, has a longer tarsus and is heavier than 
other populations. Only tail measurements seem to overlap. Further- 
more, the plumages of these two populations, though rather plain, also 
show consistent differences. Philippine A. olivaceus is overall slightly 
darker than are other populations, and although all share a contrast in 
colour between the undertail-coverts and the rest of the underparts, 
this is much more evident and striking in the paler south-eastern birds. 
The undertail-coverts, tarsal feathering and often lower belly of 
south-eastern birds are buff, typically closest to the Cinnamon-Drab 
(colour number 219c) or Tawney (38) of Smithe (1975), but sometimes 
paler (Pale Pinkish Buff; 121D) on the lower belly, whilst those of 
populations in the Philippine islands are usually a much darker russet 
colour (Cinnamon-Brown (33) to Russet (34)). Some individuals from 
the Philippines, however, have the extreme tip of the undertail-coverts 
a similar colour (Tawney (38)) to birds from the south-east. 

Comparison of vocalisations of the population of bush hens is 
dificult because few recordings exist of birds from the Philippines. 
However, four recordings of the shrieking calls (Clarke 1975) of 
Philippine birds, when compared with a series of recordings of 
shrieking calls of Australian birds, clearly sound very different 
(comparison made using the Wildlife Collection in the British Library’s 
National Sound Archive). Whilst similar in pattern, those of Australian 
birds are distinctly less harsh, higher-pitched and probably best 
described as being more melodious than those from the Philippines. 

Based on these differences in plumage and biometrics, and on 
differences in known, comparable types of vocalisations, it would seem 
reasonable to treat the populations of bush hens from the Philippine 
Islands as a species separate from those to the east, therefore 
supporting the opinion of McAllan & Bruce (1988) that there are two 
species: A. olivaceus in the Philippines, and A. moluccanus elsewhere. 
Whilst other authors have proposed the common names Rufous-tailed 
Bush Hen or Rufous-tailed Moorhen for A. moluccanus, this does not 
seem to be very appropriate. The tails of moluccanus and olivaceus are 
very similar in colour: between Prout’s Brown (121A) and Brussels 
Brown (121B). Furthermore, all the bush hens are rather plain, so that 
Plain Bush Hen is also a poor name: more appropriate alternatives are 
suggested here; Philippine Bush Hen for A. olivaceus and Pale-vented 
Bush Hen for A. moluccanus. 


Bull. B.O.C. 1998 118(2) 


71 


F. R. Lambert 


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F. R. Lambert 73 Bull. B.O.C. 1998 118(2) 


Although some authors have mentioned presence or absence of a red 
or orange swollen patch at the base of the culmen as a character 
distinguishing the different populations of bush hens, very few data are 
available concerning the actual distribution of this feature. Certainly, 
this feature can be seen on some specimens of A. moluccanus from New 
Britain and New Guinea (A. m. nigrifrons), and it is well documented 
for Australian birds (A. m. ruficrissus). However, it is not permanent, 
and is apparently associated with breeding (Marchant & Higgins 1993); 
hence its presence in other populations could have been overlooked. 
Whilst there is no present evidence that A. olivaceus can also show a 
swollen base to the culmen, this cannot be stated with certainty; nor is 
it known whether other populations of A. moluccanus, for example the 
nominate subspecies inhabiting the Moluccas, ever show this feature. 
This possible difference between the two species therefore needs 
further investigation, and indeed, if it were lacking the Moluccan 
population of A. moluccanus, might point to there being yet another 
allospecies. 

Finally, it is worth noting that Brown Crake A. akool of mainland 
Asia has long been treated as a good species, and yet it is no more 
different in appearance to bush hens from the Moluccas than are bush 
hens from the Philippines. The plumage of A. akool shows many 
similarities to A. moluccanus, with the grey of the underparts only 
slightly paler and the upperparts only slightly more olive. [The 
underparts of akool are closest to Smithe’s (1975) Light Neutral Gray 
(85); those of moluccanus are between the latter and Medium Neutral 
Gray (84), whilst olivaceus has underparts between Medium Neutral 
Grey (84) and Dark Neutral Gray (85)]. In plumage A. akool only 
differs from A. moluccanus in having a white throat (like immature 
A. moluccanus) and in lacking the buff undertail-coverts. Biometrics are 
also quite similar, though A. akool has a narrower bill (Table 1). Of 
additional interest is the fact that the underparts of A. isabellina of 
Sulawesi are almost exactly the same colour as the undertail-covert 
colour of A. moluccanus. In this suite of species, therefore, there are 
only very subtle differences in the few colours exhibited, and in the 
extent or distribution of such colours. Biometric differences, however, 
are clearly evident, and, in the case of A. moluccanus and A. olivaceus, 
combination with the subtle plumage changes provide compelling 
reasons to treat these taxa as good species. 


A new species of bush hen 


Having concluded that the Philippine, and Moluccan to Australasian 
forms of bush hen represent two good species, the question then arises 
as to how to treat the newly discovered taxon on Karakelong. This is 
apparently the only island where two bush hens are sympatric, and 
lying between the Philippines and Moluccas, one might anticipate that 
A. olivaceus and A. moluccanus would meet here. One of the species, 
based on the relatively small size and plumage of the birds seen that had 
been trapped, was identified as A. moluccanus: this being also known 
from the nearest island to the south—Sangihe—and directly to the east 


F. R. Lambert 74 Bull. B.O.C. 1998 118(2) 


on Morotoi and Halmahera (White & Bruce 1986). A. olivaceus occurs 
to the north of Talaud, on Mindanao. However, comparison with the 
specimen of the second Amaurornis from Talaud clearly shows it to be 
different to both of these species. It is therefore concluded, for reasons 
given below, and using the biological species concept, that the largest 
Amaurornis taxon from Talaud, which is sympatric with A. moluccanus, 
should also be considered a separate species. The proposed name for 
this new species of bush hen is: 


TALAUD BUSH HEN Amaurornis magnirostris sp. nov. 


Holotype 

Museum Zoologicum Bogoriense, Bogor, Indonesia, specimen 
number MZB 30.272: collected by Frank R. Lambert on 15 August 
1996; unsexed. The bird was purchased in Beo, Karakelong Island, 
Talaud, from a hunter who had trapped it near Rainis (4°14'N 
126°51’E) at an altitude of less than 100 m a.s.l. (Plate 2). 


Diagnosis 

Most similar to Amaurornis olivaceus but also superficially similar to 
A. moluccanus, which is sympatric with A. magnirostris on 'Talaud. 
Amaurornis magnirostris differs from both these species in being darker 
below with no contrasting paler-coloured undertail-coverts and in 
having a considerably bigger skull and longer, broader bill (Table 1) 
with a distinctly arched culmen. 

The skull is 8.0% wider (25.5 mm in magnirostris, measured at the 
widest point) than the largest intact skull of nine A. olivaceus 
(20.3-23.7 mm) in BMNH. Bill of A. olivaceus (measured from the 
gape to the tip) ranged from 32.2—39.4 mm in the series of 15 birds with 
intact bills in BMNH. In comparison, the bill length of the holotype of 
A. moluccanus is 45 mm, or c. 14% longer than the longest bill of A. 
olivaceus. ‘The longest-billed specimens of A. moluccanus in BMNH 
have bill lengths of 35.2 mm. 

Although the depth of the bill of A. magnirostris falls into the range 
for that of A. olivaceus when measured at its base, the depth of the 
former is 15.6% larger at the nares than that for the largest 
measurement for any of the specimens of A. olivaceus. This clearly 
demonstrates the difference in shape of the bill, which is strongly 
arched in the former species, and broad along a _ considerable 
length—that of A. olivaceus is shorter and more tapered. The large head 
and bill size of A. magnirostris can also be discerned by comparing 
measurements with A. zsabellinus, which has usually been considered to 
be a large species. 

Compared with A. olivaceus, which is most similar in size, 
measurements of the tarsus and foot suggest that, although of similar 
length, lying at the upper range limit of the latter, the leg of A. 
magnirostris 1s stouter and the feet heavier. 

A. tsabellinus is similar in size to A. magnirostris but 1s 
smaller-headed with a smaller bill, and has more slender, longer legs. 
The culmen is concave or straight, not arched as in A. magnirostris. 


F. R. Lambert 75 Bull. B.O.C. 1998 118(2) 


It should be noted that Ripley’s (1977) measurement of A. olivaceus 
and A. moluccanus (Table 2), based on an unknown sample size, differ 
in range from those provided in Table 1. Those of Parkes (1971), 
however, are in general agreement with those made by myself and 
presented in Table 1. 

Although not all vocalisations are well documented, those of A. 
magnirostris that were recorded (the “piping” calls—see under 
Vocalisations) appear to be significantly different from those of A. 
moluccanus, which are well documented in Australia (see Figure 2). 

Finally, A. magnirostris seems to be an adaptable forest bird, 
penetrating tall primary forest far from secondary vegetation, rather 
than a bird of disturbed habitats. 


Description of holotype 

Plumage very worn, with heavy body, wing and tail moult. Worn 
feathers different in colour to those that are fresh. T'ail feathers missing, 
apparently lost during capture. Presumed to be an adult (immatures of 
A. olivaceus are pale, not dark). The following description uses colour 
names and codes of swatches from the Naturalist’s Colour Guide where 
it was possible: the names from Smithe (1975) start in uppercase. 

Forehead to nape Dusky Brown (19) with paler tinge to some 
feathers; lores and feathers of orbital region similar but more sooty. 
Mantle Fuscous (21), Cinnamon-Brown (33) and. becoming very 
slightly olive on back; lower back and rump Burnt Umber (22). 
Upperwing coverts like mantle but with patches of Cinnamon-Brown 
(23) (these appearing mostly on worn feathers) more pronounced on 
greater and median coverts. Primaries and secondaries Fuscous (21) 
with diffuse Cinnamon-Brown (33) leading edges. Underwing coverts 
dark brown. Chin, throat and neck Dark Neutral Gray (83); breast and 
belly a patchy mixture of Dark Neutral Gray (83) and Fuscous (21), 
becoming more Burnt Umber (22) on flanks and feathers of tarsus. 
Undertail-coverts Dark Neutral Gray (83), not contrasting with belly. 
Iris bright red; bill long, broad and arched, the lower mandible pale 
green becoming pale blue-green on distal third; upper mandible below 
nostril and basally pale green, culmen dark olive from base to past 
nares, but distal third pale green. Legs dark olive brown with bright 
yellow frontal part to upper leg from feathering to just below ankle 
joint (1.e. covering the entire tibiotarsus). 

Biometrics are provided in Table 1. Primary three (counted 
descendently) missing, and wings frayed, but wing length estimated to 
be approximately, or slightly longer than, 168 mm (although in heavy 
moult, the absence of primary three may have resulted from 
capture/transportation). 


Etymology 

This species has been named in recognition of its large bill. The 
three taxa of “‘plain bush hens’”’ that occur from Philippines to the New 
Guinea region are similar in many respects, but the bill and large skull 
of Amaurornis magnirostris, as well as its uniform underparts, set it 
apart from the others. The large bill is its most striking character in the 


Bull. B.O.C. 1998 118(2) 


76 


F. R. Lambert 


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field. The common name of the species, Talaud Bush Hen, is chosen to 
emphasize the known range of the species, which may be very 
restricted. 


Vocalisations 

Bush hens were heard on Karakelong on many occasions, although 
never in the interior of primary forest, despite the presence of A. 
magnirostris in this habitat. The distinctive growling shrieks of bush 
hens are well known, and are made by both A. moluccanus and A. 
olivaceus. They sound different, the latter sounding harsher, when tape 
recordings are compared, but there is considerable variation in such 
calls (Beruldsen 1975). These calls are often made in antiphonal duet 
(Clarke 1975). It was not proved whether or not A. magnirostris also 
makes these “‘shrieking’’ calls, although quiet shrieks (perhaps better 
described as croaking with rasping—R. Ranft pers. comm.) can be 
heard in the background of a tape recording of the “piping’’ call of A. 
magnirostris and seem likely to have been this species. These calls sound 
quite similar to four tape recordings of the shrieking call of A. olivaceus 
held in the British Library National Sound Archive collection, but it 
was not possible to make sonagrams of the probable A. magnirostris 
calls (since sounds produce no discernible pattern, but rather a smudge 
mixed with background noises). This tape recording made by the 
author is deposited at the British Library National Sound Archive. 

The only call definitely attributable to A. magnirostris was a 
monotonous series of very loud notes that sound rather frog-like. The 
notes can best be described as low-pitched barks. Although 
significantly different from the monotonous series of calls made by 
Australian A. moluccanus, the type of call is similar in pattern, and 
hence here referred to as the piping call based on those described and 
illustrated (using sonagrams) by Clarke (1975). He observed A. 
moluccanus feeding whilst making this call. Playback of the piping call 
of A. magnirostris elicited an immediate response from calling birds, 
which quietly but rapidly investigated the area from where the playback 
emanated. Figure 2 compares two sonagrams of piping calls of A. 
olivaceus with those of A. magnirostris. Other sonagrams of the piping 
calls of A. olivaceus are provided by Clarke (1975) and Marchant & 
Higgins (1993). 


Distribution, habitat and status 

The Talaud Bush Hen is presently known only from Karakelong 
Island, in the Talaud Archipelago. It seems likely that it must also have 
occurred on neighbouring islands, but whether it survives on them (in 
particular, Salebabu and Kabaruang) remains to be determined. 
Karakelong is the largest island (c. 600 km?) in the Talaud Archipelago 
and is still endowed with good areas of forest that occur from near sea 
level to the highest peak. Evidence suggests that the Talaud Bush Hen 
is widespread on the island. Sightings in various habitats at different 
altitudes, from rank vegetation near forest edge to dry dense primary 
forest slopes, suggest that the species is adaptable and can probably 
tolerate limited habitat modification. Karakelong still has significant 


F. R. Lambert 78 Bull. B.O.C. 1998 118(2) 


natural forest cover and a diversity and abundance of wetland habitats, 
including pristine rivers, forested swamps and rank scrub and 
grasslands bordering forest. At the present time, it therefore seems very 
unlikely that the ‘Talaud Bush Hen is threatened. 

Nevertheless, since the ecological requirements of the Talaud Bush 
Hen are unknown, future changes in land use on such a small island 
could conceivably alter habitats to the extent that it becomes 
threatened. Whilst there are two protected areas totalling c. 21 000 ha 
on Karakelong, there is virtually no provision to protect these areas on 
the ground (Sujatnika et al. 1995, Sujatnika in litt. 1997; pers. obs). 
Both areas are under pressure from smallholder encroachment, and 
forest clearance for transmigration may also pose a threat in the long 
term (Whitten et al. 1987, Lambert 1997). 

Bush hens are occasionally trapped for food using snares, or caught 
by dogs. However, the trapping of terrestrial birds for food did not 
seem to be an intensive activity, and usually occurs only in the vicinity 
of villages (these being virtually confined to the coast). Hunting is 
unlikely to constitute a significant threat to the species at present levels. 

I noted that large rats (Rattus sp.) were abundant on the nearby 
island of Sangihe in 1996, particularly in plantations and the more 
degraded forest areas, but I did not encounter them on Karakelong. 
Rats (of unidentified species) are present, however, in altered habitats 
on the island (J. Wardall, zm litt. 1998) and more extensive habitat 
modification there could conceivably alter patterns of abundance. 

Studies of bones excavated on Pacific islands suggests that large 
numbers of birds that inhabited remote islands became extinct 
subsequent to the arrival of man. These studies suggest that, 
considering all bird groups known to be present on these islands, rails 
have lost most species to human impact (Steadman 1995). Whilst the 
exact cause of such extinctions will never be known, it is likely that rats 
played a role in many cases (see, for example, Atkinson 1985). If 
susceptibility to depredation by rats is accepted as a characteristic of 
island rail populations, the Talaud Bush Hen should be considered as a 
Near-Threatened species (N. J. Collar, pers. comm.). The potential 
threat from rats and other introduced mammalian predators to the 
rallids of Talaud should therefore be carefully monitored. 


Discussion 


Although the Talaud Islands are a remote archipelago, it is surprising 
that previous ornithologists who have made observations or collected 
specimens on the islands have failed to find either of the bush hens that 
live there, since both are very vocal. The Talaud Islands lie within the 
Sangihe—Talaud Endemic Bird Area (ICBP 1992, Sujatnika et al. 1995, 
Stattersfield et al. 1998), but prior to the discovery of A. magnirostris 
and the recently described Talaud Rail Gymnocrex talaudensis (Lambert 
1998), no undisputed species were known to be confined to ‘Talaud, 
although one taxon, Obscure Kingfisher, has been variously treated as a 
full species Todiramphus (Halcyon) enigma (e.g. White & Bruce 1986) 
or as a distinct subspecies of Collared Kingfisher T. chloris (e.g. Fry 


F. R. Lambert 9 Bull. B.O.C. 1998 118(2) 


et al. 1992, Inskipp et al. 1996). The discovery of two rails possibly 
endemic to Talaud is therefore of great importance, considerably 
elevating the conservation value of the islands. 

The sympatry of A. magnirostris and A. moluccanus on 'Yalaud is of 
great interest since this is the only known locality where two apparently 
ecologically similar species of Amaurornis coexist (Ripley 1977). Only 
the strikingly different, and migratory, White-breasted Waterhen A. 
phoenicurus is broadly sympatric with congeners throughout its range. 
Bush hens have not, apparently, colonised Sulawesi, where the endemic 
Isabelline Waterhen A. izsabellinus is found in many habitats from sea 
level to 1,400 m (White & Bruce 1986, Coates et al. 1997). 

Amaurornis moluccanus is not known to make regular movements and 
must be assumed to be resident on Talaud. Its presence on many 
islands from Australasia to Wallacea, however, is indicative of good 
powers of dispersal and it may be a relatively recent arrival in the 
archipelago. This could partly explain why it has not been previously 
recorded from Karakelong. That A. moluccanus has not colonised the 
Philippines might be a reflection of closer ecological requirements of 
A. moluccanus and A. olivaceus. 

On Talaud, both A. magnirostris and A. moluccanus occur in open, 
scrubby habitats, but based on the observations of A. magnirostris to 
date, it seems probable that this species is more of a forest bird, and 
that it wanders into adjacent scrub, rank vegetation and overgrown 
plantations. Although A. moluccanus may also occur in forest, this may 
be a marginal habitat for the species. 

Examination of the collection of Amaurornis bush hens in the 
BMNH clearly supports the idea that previously recognised taxa of 
these birds belong in two separate species differing significantly in 
appearance and size. That their taxonomic status has remained in doubt 
for so long is merely because careful examination of museum specimens 
was not carried out. There are clearly many unanswered taxonomic 
questions in the Oriental region (see, for example, the excellent 
annotations in Inskipp et al. 1996), and the alarming ongoing lack of 
professional taxonomists who are able to devote time to answering such 
questions should be a cause for considerable concern. Recent 
opportunistic investigations into various bird taxa (e.g. Collar & 
Andrew 1987, Brooks et al. 1992, Collar & Long 1996, Lambert & 
Woodcock 1996) suggest that avian biodiversity within this region is 
presently underestimated, and that many taxonomic revisions are 
needed. 


Summary 


This paper reviews the taxonomy of bush hens occurring from the 
Philippines through Indonesia to Australia and the New Guinea region 
and describes a new species of Rail, the Talaud Bush Hen Amaurornis 
magnirostris. 

Examination of bush hen specimens in BMNH showed that there 
were consistent differences in plumage and biometrics, and in 
comparable types of vocalisations between the Philippine population 


F. R. Lambert 80 Bull. B.O.C. 1998 118(2) 


and the populations to the east, therefore supporting the opinion that 
there are two species: A. olivaceus in the Philippines, and A. moluccanus 
elsewhere. ‘Iwo species of bush hen were found on Karakelong Island, 
in the Talaud Islands of Indonesia. One of these species was identified 
as A. moluccanus, whilst the other, collected on 6 September 1996, is 
shown to be a new species, the T'alaud Bush Hen A. magnirostris. The 
sympatry of A. magnirostris and A. moluccanus on Talaud is of great 
interest since this is the only known locality where two apparently 
ecologically similar species of Amaurornis co-exist. 'Talaud Bush Hen 
differs from the other bush hen species in having darker underparts 
with no contrasting paler-coloured undertail-coverts and in _ its 
considerably bigger skull and longer, broader bill with a distinctly 
arched culmen. Its habitat preferences may also differ, being primarily 
a forest species and occurring in primary forest far from disturbed 
areas. Vocalisations apparently differ from comparable calls of A. 
moluccanus, although it was not possible to compare these with similar 
calls of A. olivaceus. Prior to the discovery of A. magnirostris and the 
recently described Talaud Rail Gymnocrex talaudensis, no undisputed 
species were known to be confined to 'Talaud. The existence of two rails 
possibly endemic to ‘Talaud therefore considerably elevates the 
conservation value of the islands. Evidence suggests that the Talaud 
Bush Hen is widespread on the island, where significant areas of 
suitable habitat still occur. The species is therefore probably not 
immediately threatened. Nevertheless, since its exact ecological 
requirements remain unknown, and in view of the proven vulnerability 
of rails on islands, many of which have become extinct in the past, the 
‘Talaud Rail should be considered to be Near-Threatened. 


Acknowledgements 


Tim Inskipp, Nigel Redman, Adrian Long and Nigel Collar provided me with various 
literature and information unavailable in Thailand, for which I am most grateful. Barry 
Taylor very kindly provided me with draft species accounts for the Amaurornis species 
from his forthcoming book on rails of the world, and shared his views on the taxonomic 
status of bush hens. Robert Prys-Jones and Michael Walters gave me permission to 
examine bird skins at the British Museum of Natural History (Tring). I am very grateful 
to Richard Ranft and Paul Duck, who were extremely helpful in enabling me to access the 
tape recordings of calls in the British Library National Sound Archive and assisted me in 
making sonagrams. Christian Mamengko, Cober and Spener worked on the field project 
in Talaud and alerted the author to the rails on sale in Beo market, including the 
specimen that was eventually purchased and collected. Dr Soetikno Wirjoatmodjo and 
Ibu Dewi of the Indonesian Institute of Science (LIPI) provided me with assistance in 
borrowing the specimen from the Bogor Museum, and I am grateful to Dr M. Amir for 
facilitating this. Nivesh Nadee of the Thailand Institute of Scientific and Technological 
Research skilfully saved the specimen from decomposition. Richard Grimmett kindly 
compared the plumage of the specimen of A. magnirosiris in Bogor with a colour chart 
loaned by the Smithsonian Institution with the assistance of Pamela Rasmussen. Nigel 
Collar, Tim Inskipp, William Duckworth and Barry Taylor provided constructive 
comments on drafts of the manuscript. 


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Beehler, B. M., Pratt, T. K. & Zimmerman, D. A. 1986. Birds of New Guinea. Princeton 
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Beruldsen, G. R. 1975. The bush-hen in south-eastern Queensland. Australian Bird 
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Brooks, “f.. M., Evans, T. D.,. Dutson, G..'C.b;,"AndersonsG: QwiA., Asane;D. C., 
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Buckingham, R. & Jackson, L. 1987. A field guide to Australian bird song. Cassette 3. 
Bird Observers’ Club, Victoria. 

Clarke, J. H. 1975. Observations on the Bush-hen at Camp Mountain, south-east 
Queensland. Sunbird 6: 15-21. 

Coates, B. J., Bishop, D. B. & Gardner, D. 1997. A Guide to the Birds of Wallacea: 
Sulawesi, the Moluccas and Lesser Sunda Islands, Indonesia. Dove Publications, 
Alderley, Australia. 

Collar, N. J. & Long, A. J. 1996. Taxonomy and names of Carpococcyx cuckoos from the 
Greater Sundas. Forktail 11: 135-150. 

Collar, N. J. & Andrew, P. 1987. Red data birds: the cochoas. World Birdwatch 9: 5. 

del Hoyo, J., Elliott, A. & Sargatal, J. (eds). 1996. Handbook of the Birds of the World. 
Vol. 3 Hoatzin to Auks. Lynx Ediciones, Barcelona. 

Dickinson, E. C., Kennedy, R. S. & Parkes, K. C. 1991. The Birds of the Philippines. 
British Ornithologists’ Union, Tring (Check-list No. 12). 

Fry, C. H., Fry, K. & Harris, A. 1992. Kingfishers, bee-eaters and rollers. Black, London. 

Gilliard, E. & LeCroy, M. 1967. Results of the 1958-1959 Gilliard New Britain 
expedition. 4. An annotated list of the birds of the Whiteman Mountains, New 
Britain. Bull. Am. Mus. Nat. Hist. 135: 173-216. 

ICBP. 1992. Putting biodiversity on the map: priority areas for global conservation. ICBP, 
Cambridge. 

Inskipp, T., Lindsey, N. & Duckworth, W. 1996. An Annotated Checklist of the Birds of 
the Oriental Region. Oriental Bird Club, Sandy, U.K. 

Lambert, F. R. 1997. Field Assessment of the Conservation Status of Red-and-Blue Lory 
Eos histrio in Indonesia. IUCN Species Survival Commission, IUCN, Bangkok, 
Thailand. 

Lambert, F. R. 1998. A new species of Gymnocrex from the Talaud Islands, Indonesia. 
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Lambert, F. R. & Woodcock, M. 1996. Pittas, Broadbills and Asities. Pica Press, 
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Marchant, S. & Higgins, P. 1993. The Handbook of Australian, New Zealand and 
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Mayr, E. 1938. Birds collected during the Whitney South Seas expedition XL. Notes on 
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Indonesian biodiversity: the endemic bird area approach. BirdLife International, 
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from Sulawesi, Indonesia? Conservation Biol. 1: 42-48. 

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© British Ornithologists’ Club 1998 


Courtship behaviour, vocalizations, and 
species limits in Atthis hummingbirds 


by Kristof Zyskowski, A. Townsend Peterson & Daniel A. 
Kluza 


Received 4 February 1997 


Hummingbirds of the genus Atthis were first described scientifically in 
1839 by Lesson & DeLattre, based on specimens from Jalapa and 
Coatepec in central Veracruz, Mexico (A. heloisa). Some 40 years later, 
in 1878, Ridgway described a second form of the genus, A. ellioti, from 
Volcan de Fuego, in the Pacific cordillera of Guatemala. Griscom 
(1932) subsequently described another southern form A. “h.” 
selasphoroides from the highlands of Honduras, and Moore (1937) 
described another northern form A. h. margarethae from northwestern 
Mexico. 

Confusion reigned long regarding species limits in the genus. In spite 
of their original description as a separate species, populations south of 
the Isthmus of Tehuantepec were included within A. heloisa by Baird 
et al. (1874) and Boucard (1892-1895). Since then, opinions have 
differed about whether ellzot1 deserves recognition as a species, with 
some authors arguing for conspecificity (e.g. Peters 1945, Johnsgard 
1983) and others separating them as two species (e.g. Berlioz 1938, 
Friedmann et al. 1950, AOU 1983). In general, the debate seems to 
centre on the idea that the two Atthis hummingbirds differ only in 
minor details of colour; given their allopatric distributions, the decision 
hinged upon taxonomic viewpoint only. 


K. Zyskowski et al. 83 Bull. B.O.C..1998 118(2) 


Even the distinctiveness of the genus has been disputed, with 
the suggestion that it should be merged into Selasphorus (Johnsgard 
1983), in spite of its perhaps closer relationship with Stellula 
(Ridgway 1892, Wolters 1976, Sibley & Monroe 1990). The close 
relationship of the genera Archilochus, Atthis, Calypte, Stellula, and 
Selasphorus, together with other small, gorgeted hummingbirds is 
supported by skeletal synapomorphies, but little resolution of 
relationships has been possible within the clade (R. L. Zusi pers. 
comm.). Howell & Webb (1995) merged Atthis into Selasphorus, and 
combined Archilochus, Calypte, and Stellula into Archilochus with no 
comment or justification. We regard this arrangement as preliminary 
and arbitrary. 

Unfortunately, little has been published regarding the ecology, 
behaviour, and vocalizations of either form of Atthis. Especially 
relevant to the question of species limits, the displays and vocaliz- 
ations associated with courtship have been described only briefly 
by Howell & Webb (1995) and Skutch (in Bent 1940). The purpose 
of the present paper is to present descriptions of the courtship 
displays and vocalizations of the northern populations of these 
hummingbirds, compare with descriptions of these displays in southern 
populations, to point out differences in behaviour and morphology 
between the two forms, and to comment on implications for their 
specific status. 


Methods 


As part of avifaunal inventory studies, KZ and DAK observed Atthis 
hummingbirds at two localities in cloud forest in the Sierra Mazateca in 
northern Oaxaca: 29-31 January 1994, at Puerto de la Soledad, a 
microwave station at the highest point along the road from Teotitlan 
del Camino to Huautla de Jiménez, Oaxaca (specific locality: Distrito 
de Teotitlan del Camino, Puerto de la Soledad, GPS coordinates 
18°9.951'N, 96°59.891'W, 2280 m); and 21 May-—2 June 1994 near San 
Martin Caballero, a town in an isolated northeastern spur of the Sierra 
Mazateca (specific locality: Distrito de Teotitlan del Camino, 1 km NE 
San Martin Caballero, 18°6.721'N, 96°38.426’'W, 1470m). Two 
specimens were preserved as vouchers [OMVP 1041 (male) and 1130 
(female)|, deposited at the Museo de Zoologia, Facultad de Ciencias, 
Universidad Nacional Autonoma de México. A. ellioti was observed by 
ATP on 3-10 July 1995 in cloud forest in Parque Nacional Los Andes, 
Volcan Santa Ana, Departamento de Santa Ana, El Salvador. 

To assess differences in vocalizations, we studied recordings of Atthis 
hummingbirds provided by the Cornell Library of Natural Sounds, 
and compared them to vocalizations heard during our field studies. 
This material included recordings by Theodore A. Parker III of A. 
heloisa (L.NS 17214) from above Puerto Los Mazos, Jalisco, Mexico, 
and of A. ellioti from Cerro Verde, Santa Ana, El Salvador, made by 
W. A. Thurber. 

To provide a preliminary assessment of morphological variation, we 
examined specimens of both forms in the collections of the University 


K. Zyskowski et al. 84 Bull. B.O.C. 1998 118(2) 


Figure 1. Distribution of hummingbirds of the genus Atthis. Black areas indicate 
populations of A. heloisa; dark grey indicates populations of A. ellioti; and light shading 
indicates probable continuity of populations in appropriate habitats. 


of Kansas Natural History Museum, Field Museum of Natural 
History, Southwestern College, and the Museo de Zoologia, Facultad 
de Ciencias, Universidad Nacional Autonoma de México. In all, we 
inspected 41 males and 13 females of the northern populations, and 7 
males and 2 females of the southern populations. Additional specimen 
information was kindly provided by the Louisiana State University 
Museum of Natural Science. 


Distribution 


The distribution of Atthis hummingbirds generally follows the major 
mountain systems of northern Mesoamerica (Fig. 1). Populations 
assigned to A. h. heloisa range from central Tamaulipas south in the 
Sierra Madre Oriental to the Nudo de Zempoaltépetl of northern 
Oaxaca, in the interior in the vicinity of the Federal District and on 
Cerro San Felipe in northern Oaxaca, and through the Sierra Madre 
del Sur of Guerrero and Oaxaca; but the species was not detected on 
the peripheral montane forest island of Cerro Piedra Larga in 
east-central Oaxaca (Peterson et al. in prep.). An isolated population 
apparently occurs in southcentral San Luis Potosi in the vicinity of 
Alvarez; Salvin & Godman (1879-1904) reported a specimen, perhaps 
of doubtful veracity, collected by A. Dugés in Guanajuato (not 


K. Zyskowshi et al. 85 Bull. B.O.C. 1998 118(2) 


included on map for lack of a more specific locality). A specimen 
collected by Mario del Toro Avilés at ‘‘Montanas Gineta’’, Oaxaca, is 
an example of A. h. heloisa outside of that form’s range, another 
example of that collector’s notoriously unreliable labelling of specimen 
material (Binford 1989). 

Populations referred to as A. h. margarethae are restricted to the 
coastal slopes in Sinaloa, Nayarit, and Jalisco, and then apparently in 
the Transvolcanic Belt east to western Estado de México. Their 
absence from the higher peaks of the main body of the Sierra Madre 
Occidental is odd, given their occurrence in similar habitats in the 
Transvolcanic Belt. Our limited reexamination of the characters used 
by Moore (1937) indicated that the differences appear real, although the 
distributional gap that he mentioned does not, based on ranges outlined 
in Friedmann et al. (1950) and Howell & Webb (1995). Two female 
specimens described as A. morcomi by Ridgway (1898) from 
southeastern Arizona appear to represent either stragglers or 
mislabelled specimens; Bangs (1929) pointed out that both fall 
completely within the range of variation of A. h. heloisa. Humming- 
birds of this genus have not been found subsequently at the type 
locality, in spite of its extreme popularity among birdwatchers. 
Although these extralimital records might suggest seasonal or 
altitudinal movements, evidence available is insufficient to demonstrate 
this phenomenon convincingly. 


Courtship behaviour 


Observations of courtship behaviour of A. h. heloisa were as follows. 
Males were distributed relatively uniformly through the habitat, 
especially along ridgetops, frequently perching on high, exposed 
branches of Podocarpus sp. in disturbed vegetation along trails. Females 
were less obvious, often hidden nearby in dense vegetation closer to the 
ground. Individuals of both sexes were observed to feed low to the 
ground from red-flowered Salvia sp. (Lamiaceae) at Puerto de la 
Soledad, and from yellow-flowered Palicourea galeottiana (Rubiaceae) 
at San Martin Caballero. 

Males sang from perches, and appeared to be consistent in their use 
of particular branches, being seen in the same positions on as many as 
12 consecutive days. Vocalizations included a rather soft, short tsz! 
given by individuals of both sexes. Perched males, however, gave the 
same tsz!, followed by a thin whistling weeeeeeeeeew that rose and then 
fell in pitch, lasting a total of two or three seconds (Fig. 2), the whistled 
portion being reminiscent of songs of Calypte costae (Wells et al. 1978, 
KZ pers. obs.). Some immature-plumaged males at San Martin 
Caballero were heard to sing two or three repetitions of a briefer 
version of this song in quick succession, much as described by Wells 
et al. (1978) for C. costae. 

Perched males oriented themselves towards nearby females, which 
were often perched or foraging. As frequently as once per minute, a 
male would fly to within 10 cm of a female and hover horizontally in 
front of her, spreading his gorget and cocking his spread tail vertically 


K. Zyskowski et al. 86 Bull. B.O.C. 1998 118(2) 


8 


Frequency (kHz) 


0 0.5 1.0 1:5 
Time (s) 


Figure 2. Sounds made by Atthis hummingbirds: wing noise (top) and song (middle) of 
A. heloisa, recorded in Jalisco, Mexico; and song of A. ellioti (bottom), recorded in El 


Salvador. 


over his back, but was not observed to make any display dive, as do 
other related genera (Wells et al. 1978, Johnsgard 1983). During the 
hovering, the male’s wings produced a wavering thrumming noise (Fig. 
2; Robins & Heed 1951), and he often followed the female’s movements 
closely. The noise produced by the wings was similar to that of 
courting Selasphorus platycercus, although somewhat softer (KZ pers. 


K. Zyskowski et al. 87 Bull. B.O.C. 1998 118(2) 


obs.). Occasionally, while courting females, males flew in horizontal 
loops as long as 8 m, making the wing noise continuously. The wing 
noise was also noticeable when males flew in non-courtship behaviours 
such as foraging, but whether it is always produced during flight is 
unclear; Howell & Webb (1995) also noted that wing noise is louder 
during displays, but produced continuously. Immature males were not 
seen to court females; nor were immature males or females heard to 
produce wing noise when flying. Observations in January included both 
singing and courtship, but in May only singing was noted, suggesting 
that nesting was already well underway or completed. 

These observations contrast in some respects with those of Skutch 
(Bent 1940) of A. elliott in Guatemala and of Thurber et al. (1987) from 
El] Salvador. They described assemblies of males spaced 25-30 m apart, 
with no other such assemblies detected within 2 km. Similar to our 
observations, the males sang from exposed perches, but their song was 
described as rising and falling in pitch, more rich and varied (lacking 
the whistling quality) than in A. helotsa, and lasting 30-40 seconds, 
much longer than in A. heloisa, as was borne out by the recordings we 
studied (Fig. 2). No pronounced wing noise was noted. Excepting the 
latter point, these differences are largely in accord with descriptions in 
Howell & Webb (1995). Displaying males apparently moved their 
gorgets, and often sang while in looping flights, but were not observed 
to approach the females closely (but see Howell & Webb 1995). 

Hence, several marked differences seem to exist in the vocalizations 
and courtship behaviours of the two forms of Atthis hummingbirds. 
The northern form (A. heloisa) sings a simpler song and only while 
perched, approaches closely to females while in flight, and produces a 
loud humming wing noise while flying. Observations (ATP) at close 
range of A. ellioti in El] Salvador indicated that its wing noise is much 
quieter and less throbbing than in A. heloisa; this observation 
contradicts a brief mention of display behaviours in Howell & Webb 
(1995). Finally, and perhaps most interesting, is the possibility that the 
southern birds display in groups (leks?), whereas the northern birds 
show no obvious tendency towards clumping; S. N. G. Howell, 
however, reports observations of clumped and nonclumped displaying 
males in each form (pers. comm.). 


Morphology 


Our examinations of study skins revealed several differences between 
males of the northern and southern forms of Atthis. The inner web of 
the outermost primary of all adult males of A. heloisa examined was 
notched for an average of 6.5 mm from the feather tip (Fig. 3). No 
females or immature males showed this modification, nor did any 
individual examined of A. elliot. This structural modification, noted by 
Ridgway (1892), probably accounts for the humming noise produced 
by adult male A. heloisa (Monroe 1968). An interesting sidelight of this 
observation, if the pulses in the noise represent wingbeats (Fig. 2), is 
that the wingbeat frequency for A. heloisa can be calculated at 61.3 
beats per second. 


K. Zyskowshi et al. 88 Bull. B.O.C, 1998 118(2) 


Figure 3. Shape of outer primaries of left wings in Atthis elliot: (left, FMINH 42768) and 
A. heloisa (right, KU 46137), both adult males. 


This notch of the inner web of the outer primary in A. heloisa is the 
most extreme within the five closely related genera Selasphorus, Atthis, 
Archilochus, Calypte, and Stellula. The latter three genera and 
Selasphorus flammula show no notch of the outer primary, whereas 
S. platycercus shows a notch of the distal portion of the feather only. 
Other Selasphorus (S. rufus, S. sasin, and S. scintilla; S. ardens not 
determined) have a pointed outer primary, but no notch. 

‘The colour of the two Atthis forms’ gorgets differs, in that gorgets of 
A. helotsa are of a rich magenta purple or bluish purple, but those of A. 
elliott lack blue almost completely and are decidedly more reddish, 
especially in Honduran A. e. selasphoroides (Monroe 1968), even when 
specimens of similar time since collection are compared. Additionally, 
the length of the gorgets of adult males may differ, although this feature 
is difficult to evaluate quantitatively; gorgets of A. elliott seem to be 
about 3-5 mm longer than those of A. heloisa. Our measurements of 
body dimensions were based on too few individuals to permit statistical 
testing, but seem generally to support the notion that A. ellioti is 
somewhat smaller than A. heloisa in bill and tail length, but slightly 
larger in wing length, as documented by Ridgway (1892, 1911). 


Species limits 


The sum of the information presented above is that the northern and 
southern forms of Atthis differ in several regards. "The two forms differ 
in courtship behaviour, song structure, wing morphology, and 


K. Zyskowski et al. 89 Bull. B.O.C. 1998 118(2) 


coloration. Character distributions are nonoverlapping in several cases, 
and their status as valid phylogenetic species is unquestionable. 

The unresolved question, however, is whether they should be 
considered as representing two biological species. Because of their 
allopatric distributions, no test of sympatry is available to aid in this 
decision. Comparisons with sympatric species pairs in related genera 
are not illuminating because sympatry among congeners (e.g. Calypte 
spp.) is relatively rare; however, species pairs in more distantly related 
hummingbird clades (e. g. Amazilia spp.) are maintained in sympatry 
even though they are more similar in courtship behaviours than the 
Atthis species treated herein. ‘The marked differences in courtship 
behaviour and associated morphological modifications strongly suggest 
that they would not interbreed were populations to come into contact. 
Hence, we recommend that these two forms be recognized as full 
biological species. 


Acknowledgements 


This contribution is dedicated to the memory of Ted Parker, for his many insights, 
observations, and records that have enriched so much of our work with Neotropical birds, 
and for his contributions of the recordings used in this paper. We would like to thank our 
field companions, especially Griselda Escalona-Segura, Blanca E. Hernandez-Banos, and 
Laura Gonzales-Guzman, for their assistance and support. We also thank the curators 
and staft of the American Museum of Natural History; Natural History Museum, Tring; 
Canadian Museums of Nature; Carnegie Museum of Natural History; Delaware Museum 
of Natural History; Field Museum of Natural History; Louisiana State University 
Museum of Natural Science; Moore Laboratory of Zoology; Musée d’Histoire Naturelle 
de Paris; Museo de Zoologia, Facultad de Ciencias, U.N.A.M.; Museum of Comparative 
Zoology, Harvard; Museum of Vertebrate Zoology, Berkeley; Royal Ontario Museum; 
San Diego Natural History Museum; Southwestern College; Texas Cooperative Wildlife 
Collections; Universidad Michoacana San Nicolas de Hidalgo; University of Arizona; 
Western Foundation for Vertebrate Zoology; and Yale Peabody Museum for access to 
specimens and data under their care; Gary R. Graves, Steve N. G. Howell, Adolfo G. 
Navarro-Sigtenza, James V. Remsen, and Mark B. Robbins for critique of the 
manuscript; and Greg F. Budney of the Library of Natural Sounds, Cornell University, 
for providing the sound recordings. 


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Ornithol. Union, Washington, D.C. 

Baird, S. F., Brewer, T. M. & Ridgway, R. 1874. A History of North American Birds. 
Land Birds. Little, Brown & Co., Boston, Massachusetts. 

Bangs, O. 1929. Atthis heloisa morcomi Ridgway, not a valid subspecies. Condor 29: 
118-119. 

Bent, A .C. 1940. Life histories of North American cuckoos, goatsuckers, hummingbirds, 
and their allies. U.S. Nat. Mus. Bull. 176: 1-506. 

Berlioz, J. 1938. Notes critiques sur les trochilidés. Oiseau 8: 3-19. 

Binford, L. C. 1989. A distributional survey of the birds of the Mexican state of Oaxaca. 
Orn. Monogr. 43: 1-418. 

Boucard, A. 1892-1895. Genera of Hummingbirds. London. 

Friedmann, H., Griscom, L. & Moore, R. T. 1950. Distributional check-list of the birds 
of Mexico, part 1. Pacific Coast Avifauna 29: 1-202. 

Griscom, L. 1932. New birds from Honduras and Mexico. Proc. New England Zool. Club 
13: 55-62. 

Howell, S. N. G. & Webb, S. 1995. A Guide to the Birds of Mexico and Northern Central 
America. Oxford Univ. Press. 


A. Gamauf & M. Preleuthner 90 Bull. B.O.C. 1998 118(2) 


Johnsgard, P. A. 1983. The Hummingbirds of North America. Smithsonian Institution 
Press, Washington, D.C. 

Monroe, B. L., Jr. 1968. Birds of Honduras. Orn. Monogr. 7: 1-458. 

Moore, R. T. 1937. Four new birds from north-western Mexico. Proc. Biol. Soc. Wash. 
50: 95-102. 

Peters, J. L. 1945. Check-list of Birds of the World. Vol. 5. Harvard Univ. Press. 

Ridgway, R. 1892. ‘The humming birds. Rept. U.S. Nat. Mus. 1890: 253-383. 

Ridgway, R. 1898. Description of a new species of hummingbird from Arizona. Auk 15: 
325-326. 

Ridgway, R. 1911. Birds of North and Middle America, part 5. U.S. Nat. Mus. Bull. 50: 
1-859. 

Robins, C. R. & Heed, W. B. 1951. Bird notes from La Joya de Salas, Tamaulipas. 
Wilson Bull. 63: 263-270. 

Salvin, O. & Godman, F. D. 1879-1904. Biologia Centrali-Americana: Aves. Taylor and 
Francis, London. 

Sibley, C. G. & Monroe, B. L., Jr. 1990. Distribution and Taxonomy of Birds of the World. 
Yale Univ. Press. 

Thurber, W. A., Serrano, J. F., Sermeno, A. & Benitez, M. 1987. Status of uncommon 
and previously unreported birds of El Salvador. Proc. West. Found. Vert. Zool. 3: 
109-293. 

Wells, S., Bradley, R. A. & Baptista, L. F. 1978. Hybridization in Calypte 
hummingbirds. Auk 95: 537-549. 

Wolters, H. E. 1976. Die Vogelarten der Erde, Lief. 2. Paul Parey, Berlin. 


Address: Natural History Museum, The University of Kansas, Lawrence, Kansas 66045, 


© British Ornithologists’ Club 1997 


A new taxon of the Barred Honeybuzzard 
Pernis celebensis from the Philippines 


by Anita Gamauf & Monika Preleuthner 
Received 2 Fuly 1997 


The Barred Honeybuzzard Pernis celebensis is restricted to Sulawesi 
and the Philippines. Two subspecies have been distinguished: The very 
colourful and uncrested nominate subspecies Pernis celebensis celebensis 
Wallace, 1868, is found on Sulawesi, including Muna and Peleng 
Islands; the paler banded and crested Pernis celebensis steere1 Sclater, 
1919, has been reported from all over the Philippines, except Palawan 
(Delacour & Mayr 1946, Brown & Amadon 1969, del Hoyo e¢ al. 1994). 
Dickinson et al. (1991) compiled a list of 17 islands where the 
occurrence of this subspecies has been documented. 

In the course of an ecomorphological study of Philippine birds of 
prey (Gamauf et al. 1998) we carried out morphological measurements 
on 21 raptor species in various museum collections. Sixty external 
measurements were taken from each specimen. While comparing the 
specimens from different Philippine islands, we were struck by clearcut 
differences in colour and plumage pattern among birds from northern 
and southern provenances. To investigate these geographic differences 
in greater detail, we looked for representatives of this species in 30 


A. Gamauf & M. Preleuthner 91 Bull. B.O.C. 1998 118(2) 


different museum collections. Finally, from nine museums (for 
abbreviations of museum names, see Acknowledgements), a total of 37 
specimens was available, from 10 different islands. In addition, 75 
observations in the field were available, carried out over a period of 
more than 9.5 months. 


Plumage variation 

The most striking difference between the two population groups is 
the uniformly brown colour of the adults in the northern population 
which does not display the rich contrast and coloration of specimens 
from the southern islands. This may be the reason for some confusion 
in the past concerning age classes, since plumage characteristics were 
often used to determine age (Stresemann 1940, Brown & Amadon 
1969). The holotype of steerez (Sclater 1919), now in the Natural 
History Museum, Tring—BM 1896.4.15.40, is an adult male of the 
south Philippine subspecies which was collected by Steere on 17 
February 1888, in San Antonio (Negros). We agree with Sclater’s 
statement that “. other examples from Samar, Mindanao, and 
Basilan closely resemble the type ...’’, since we were able to confirm 
the occurrence of representatives of the southern population on those 
islands. 


Morphological variation 

Table 1 gives a comparison of 14 external morphological 
measurements of individuals from northern and southern provenances. 
From the total of 37 specimens we could include 29 sexed and fully 
feathered birds in a discriminant function analysis (12 from the north 
and 17 from the south). With a combination of 6 variables (Fig. 1) we 
were able to discriminate unambiguously between populations as well 
as between age classes. In the northern population the separation 
according to sex and age class was clearcut without any overlap: females 
are larger than males, immatures are smaller than older birds (adults 
and subadults) in some measurements. In the slightly smaller southern 
form no clear discrimination was found between the sexes. This may be 
partly due to incorrect sexing of the museum specimens, as has been 
proven for other species with much more pronounced sexual 
dimorphism. Nevertheless, the age class could be determined correctly. 
Discriminant function (DF) 1 concerns characters related to the mode 
of handling the prey as well as the flight apparatus. It segregates the 
subspecies largely by the length of the bill and middle toe. A negative 
correlation exists with the number of notches and Kipp’s distance. 
Along DF 2 the honeybuzzards fall into two distinct groups largely 
according to the length of the tail as a character for flight (lift and 
ability for manoeuvring) and the tarsus length (presumably connected 
with feeding habits). 

Thus, based on the characters discussed above, the northern 
population is distinct in both plumage pattern and morphology. Every 
specimen can be clearly diagnosed. We therefore consider this 
population to represent a third taxon, for which we propose the 
name 


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Figure 1. Separation of the two Philippine populations of the Barred Honeybuzzard 
Pernis celebensis (12 study skins from the north, filled symbols; 17 study skins from the 
south, open symbols) according to discriminant function analysis of 6 morphological 
variables (bill length, length of the middle toe, number of notches, Kipp’s distance, 
length of the tail, tarsus length). Immatures are marked by round symbols. 


Pernis celebensis winkleri subsp. nov. 


Holotype. Adult male, from Bataan, Luzon, collected by O. Koch, 17 
August 1881, Zoologisches Museum der Humboldt Universitat Berlin, 
Germany, cat. no. ZMB 25.464 (Fig. 2). This is the specimen listed as 
“immature?” by Stresemann (1940, pp. 192/193). 

Diagnosis. ‘The subspecies can be distinguished unequivocally in 
subadult and adult specimens. In contrast to winkleri, individuals of 
steeret are much more contrasting in plumage. The ground colour of 
crown and neck is paler with dark stripes, the long pointed crest (up to 
73 mm) is black. The throat is white with black mesial and lateral 
stripes. The breast is whitish to buffy with bold black streaks. The 
lower breast is white with narrow rufous-brown bars. Lower belly, 
undertail coverts, leg feathers and underwing coverts are barred 
medium to dark brown and white. All illustrations in publications to 
date show steerez (duPont 1971, Brown & Amadon 1969, Weick 1980, 


A. Gamauf & M. Preleuthner 95 Bull. B.O.C. 1998 118(2) 


Figure 2. A. Holotype of Pernis celebensis winkleri subsp. nov. (Zoologisches Museum 
der Humboldt Universitat Berlin, Germany, cat. no. ZMB 25.464). B. Typical adult 
representative of Pernis celebensis steerei, Universitets Zoologiske Museum, Kobenhavn, 
Denmark, cat. no. 940). 


del Hoyo et al. 1994). In the course of our investigations we discovered 
several misidentified specimens among study skins, as was also noted 
by Dickinson et al. (1989). These obvious errors are due to close 
similarities between corresponding subspecies of the Philippine 
Hawk-Eagle Spizaetus philippensis (Preleuthner & Gamauf 1998) and of 
the Barred Honeybuzzard Pernis celebensis. The respective northern 
subspecies from both species resemble each other, as also is the case 
with the southern subspecies. Whether this could be caused by mimicry 
will be discussed elsewhere. 


A. Gamauf & M. Preleuthner 96 Bull. B.O.C. 1998 118(2) 


Description of holotype. Generally a dark uniformly brownish 
coloured bird. Especially as the holotype was originally a mounted speci- 
men, the slender honeybuzzard form can be easily recognised. Crown and 
the sides of the head are hazel to blackish-brown with fine black streaks on 
the sides and with broader black streaks on the crown. The short bluish- 
grey imbricated feathers extending from the lores to the supercilium and 
the anterior part of the auriculars, together with additional morphological 
measurements, suggest that the specimen may be sexed as male. The 
longest lanceolate feather of its pointed crest has a length of 57 mm. The 
throat is buffy white and every feather has a black shaft. Additionally, the 
throat is divided by a black median stripe and bordered by black lateral 
moustachial stripes. ‘The breast feathers are hazel to cinnamon brown 
with more or less bold black streaks. Belly and undertail coverts are 
uniform cinnamon brown like the underwing coverts and the legs. The 
plumage of the anterior part of the legs covers the upper third of the 
short tarsus. The primaries are barred on their distal part, the proximal 
part is indistinctly mottled. The wing-shape 1s typical for a forest-living 
honeybuzzard, relatively short and rounded, and the wing tips extend 
more than halfway to the tail tip. ‘The upperparts are uniform dark 
umber to sepia brown, with narrow paler edges; the primaries are 
blackish-brown. The long tail is the same ground colour on the dorsal 
side, the broad black subterminal bar separated by a broader pale 
greyish-brown unmarked zone from 3—4 narrower bars basally. The 
cere is dark grey and the bill black, only the innermost half of the lower 
mandible is warm buff. The legs are a deep yellow. 


Etymology.We have chosen to name this subspecies after the 
well-known ornithologist Prof. Dr Hans Winkler, director of the 
Konrad Lorenz-Institute for Comparative Ethology, Austrian 
Academy of Sciences, and expert in the field of ecoethology. We wish to 
emphasise our appreciation of his scientific competence and our 
gratitude for his patience and encouragement of our work. 


Paratypes. Eight specimens of winkler1 from Luzon are herein 
designated paratypes (1 adult male and 4 subadult/adult females, 2 
immature males and 1 immature female). Adult male, 05.06.1894, 
Molino, Isabela Province; BMNH, cat. no. 394.6.14.13. Adult female, 
15.03.1894, La Trinnadad, Benguet Sub-province, Mountain Province; 
BMNH, cat. no. 334.99.14.12. Adult female, 27.07.1894, Manito, 
Albay Province; BMNH, cat. no. 399.97.5-13.143. Subadult female, 
01.08.1959, Pangil, N.A. Icarangal, Laguna Province; AMNH, cat. 
no. 782423. Subadult female, 01.12.1969, Ipo Dam, Bulacan Province; 
DMNH,, cat. no. 3622. Immature male, 02.07.1958, Bataan Province, 
J. E. duPont; DMNH, cat. no. 1300. Immature female, Saray, Pakil, 
Laguna Province; DMNH, cat. no. 43724. Immature male, 30.03.1975, 
Diman, Pangil, Laguna Province; CMNH, cat. no. 47656. Seven 
additional study skins of the northern population were available from 
Polillo (1 immature, AMNH), Catanduanes Island (1 unsexed 
subadult, DMNH; 1 immature male, PNM; 1 juvenile female, PNM), 
Marinduque Island (1 subadult male, DMNH), and Sibuyan Island (1 


A. Gamauj & M. Preleuthner 97 Bull. B.O.C. 1998 118(2) 


subadult male, DMNH). One adult female, without designated locality 
(BMNH, cat. no. 1057), belongs also the subspecies winklert. 
Plumages. Adult: All adult birds are dark and with little contrast. The 
crown and the side of the head are buffy to blackish brown with heavy 
streaks broadening towards the crown. Winkleri is characterised by a 
prominent pointed crest (up to 60 mm). Adult males show bluish-grey 
feathers at the side of the head as described for the holotype. In adult 
females, the bluish-grey zone covers only the lores; but this 
characteristic appears also in the other two subspecies. In comparison 
to Pernis apivorus and Pernis ptilorhyncus this zone is clearly smaller, 
especially in males. The colour of the iris is bright yellow, cere and bill 
are dark grey to black. The throat is white to buffy white, often with 
fine black longitudinal stripes along the shafts, divided by a black 
median stripe and bordered by black lateral moustachial stripes. Breast 
feathers are chestnut to hazel with more or less bold black streaks. 
Lower breast and belly are uniformly chestnut brown. The feathered 
tibiotarsus and the undertail coverts are cinnamon. The underwing 
coverts are either uniformly coloured or of a mottled brown showing no 
bars whatsoever. The upperparts including the tail are uniformly 
coloured, umber to sepia with somewhat darker primaries. Freshly 
acquired wing and tail feathers are tipped with white. The long tail 
feathers have a broad black subterminal bar adjacent to a broader 
unmarked zone and usually 5—6, occasionally up to 7, narrower dark 
bars basally. The innermost bars are narrow and closely spaced. They 
are barely visible in perching birds. The full coloration is acquired 
approximately at the age of 3 years, as is the case with P. apivorus and 
probably also P. ptilorhyncus. Second year plumage: The feathers at the 
crown and the neck are paler than in adults. A tawny mask is clearly 
visible. The typical bluish-grey feathered zone is already developed. 
The characteristic throat pattern with the black mesial stripe and the 
lateral moustachial stripes is less contrasting and pronounced than in 
the adult. The breast is white to buffy with more or less distinct streaks. 
Lower belly, legs and undertail coverts are usually darker. The 
underwing coverts are buffy to washed brownish without pattern. Only 
in one specimen chestnut axillary patches could be observed. The pale 
band on the upperwing (present in the first-year plumage) becomes 
darker, smaller and also more indistinct. Pattern and colouring of 
primaries, secondaries and tail feathers generally resemble the adult 
plumage. Immature: The first-year ventral plumage is white (only in 
one specimen buffy) and there is almost no variation, unlike the 
immatures of P. apivorus or P. ptilorhyncus orientalis. The head is 
white, neck and crown are whitish to buffy with fine dark streaks. The 
long pointed crest is black and clearly visible. A blackish mask in the 
form of a dark eyeline reaches from the bill to the auriculars. The cere 
is yellow, the bill blackish-grey and the inner half of the lower 
mandible warm buff to yellowish. On the ventral side the distal half of 
the primaries is blackish-brown, with up to 8 bars on the longest 
primary (usually the 6th). On the proximal half the bars are reduced; 
the wing-base, therefore, appears almost white. The secondaries are 
greyish-brown with bars less contrasting than in the adult plumage. 


A. Gamauf & M. Preleuthner 98 Bull. B.O.C. 1998 118(2) 


The back is sepia brown, and the uppertail coverts vary from cinnamon 
to buffy and whitish. Especially the greater and median wing coverts 
show wide pale edgings, which were, however, lacking in a few newly 
fledged birds. ‘The secondaries are of a deep sepia colour, the primaries 
blackish brown. The tail feathers are sepia with 6-8 regularly spaced 
bars, in some individuals the areas between the bands becoming 
increasingly more closely set towards the tail-base. Feather tips are 
white and more pointed than in adults. The feet are yellow. 
Presumably, based on his knowledge of the European Honeybuzzard 
P. apivorus, Stresemann (1940) assumed that the specimen here 
designated the holotype of winkler1 might be immature. This mis- 
interpretation was due to the fact that most juveniles of P. apivorus are 
almost exclusively brown on the ventral side and thus resemble the 
adults of winklert. 

Specimens examined. Including the holotype, 16 study skins of 
winklert were available from 5 different museum collections (see above), 
for steerer 21 study skins were examined at 7 collections (BMNH 5, 
UPLB 4, PNM 3, USNM 3, DMNH 3, AMNH 2, UMZC 1). In the 
course of our study winklert has not been observed in the field, whereas 
75 observations of steerez were recorded, all on Mindanao. Altogether 
this species could be observed for 7.41 hours between January and 
April 1993, November 1993 to February 1994, and March to July 1994. 

Distribution. The occurrence of winkleri on Luzon (9), Polillo (1), 
Catanduanes (3), Marinduque (1), and Sibuyan (1) is documented by 
the study skins mentioned above. For steerei, specimens were available 
from Samar (5), Negros (1), Siargao (1), Mindanao (11), and Basilan 
(3). According to these data, in the east the borderline between the two 
subspecies runs along the Bernardino Strait between Luzon and Samar. 
‘Towards the west it follows the Sibuyan Sea south of Romblon. Since 
no study skins from Mindoro could be found during our investigations, 
the division line remains uncertain for the western part of the 
Philippine Archipelago. According to literature compiled by Dickinson 
et al. (1991) P. celebensis has been documented for seven additional 
islands. From our distribution data it can be concluded that six of those 
islands (Leyte, Tablas, Bohol, Tawitawi, Dinagat, Masbate) are 
inhabited by steerez, whereas Romblon, because of the close vicinity to 
Sibuyan, may probably be inhabited by winkleri. The present 
borderline between the two subspecies can be interpreted as the result 
of changes in the sea level during the Pleistocene (Diamond & Gilpin 
1983, Heaney 1985), as discussed in Preleuthner & Gamauf (1998). 

Habitat and conservation. Vhe Barred Honeybuzzard is morphologi- 
cally adapted to tropical rainforests (Gamauf et al. 1998). More than 
half of the 75 observations (steerez) have been made in lowland primary 
rainforests or secondary forests selectively logged 2—3 decades ago. The 
habitat choice of lowland areas is confirmed by our findings in the field 
as well as by the label data of the skins examined. Half of the 
observations were made at altitudes between 90 and 200m a.s.l., the 
rest more or less evenly distributed up to 1400 m a.s.1. (highest location 
1550 m a.s.].). Of 11 skins bearing altitudinal data, 9 originate from 
areas between sea level and 400 m a.s.]. Another specimen of steerez was 


A. Gamauf & M. Preleuthner 99 Bull. B.O.C. 1998 118(2) 


collected between 610 and 760m a.s.l. (Goodman & Gonzales 1990). 
As has been shown by Collins et al. (1991), the condition especially of 
the lowland forests is rather poor. ‘The major part of the rainforest has 
been lost due to logging and Raingin (shifting cultivation) activities. In 
1988 only about 17 700 km? of intact lowland forest still existed. The 
recent extent of the rainforest is not exactly known, but considering the 
prior deforestation rate the present range of the potential habitat for 
winkleri can be estimated at 5000 km/?, for steerei at 7500 km? only 
(Preleuthner & Gamauf 1998). While according to Hauge et al. (1986) 
the Philippines ranked high among tropical countries in the rate of 
deforestation, as well as in the extent of deforested area in the 1980s, 
some years later Collins et al. (1991) classified the Philippines as 
arguably the worst case in tropical Asia. 

Steere: was found at four study sites on Mindanao (NW and E Mt. 
Kitanglad area/Bukidnon, Carmen-Cantilan and PRI, former PICOP/ 
Surigao del Sur). For these four areas population densities were 
estimated. In the lowland forests of PRI (58 km’, 90-180 m a.s.1.) we 
found 7-8 pairs Sieh 14/100 km?) and in Carmen- Cantilan (27 km? 
80-540 m a.s.l.) 2-3 pairs (7—11/100 km? ). The density at higher 
elevations was up to 2.6 times lower than in these lowland areas. At 
NW Mt. Kitanglad (590-1400 m a.s.l.) 3-4 pairs were found in 45 km? 
(7-9 pairs/100 km’), and on the eastern side of the same massif 
(900-1800 m a.s.1.) 2 pairs were resident in the lower part of the 38 km? 
study site (5 pairs/100km7). A population density of 8.3-10 
pairs/100 km* was taken as a basis for estimating the total number for 
both populations, because winkleri could not be observed in the field. 
According to these calculations 400-500 pairs of winklert may still live 
in the potential habitat of 5000 km’. Using the same approach for 
steere1 the corresponding number is 600-750 pairs in an area of 
7500 km? of potential habitat. However, the actual number of winkleri 
may be much smaller since Danielsen et al. (1993) have classified the 
Barred Honeybuzzard as scarce for the Sierra Madre region, Luzon, 
and steere: could not be found at Mindoro, Negros and Sibuyan in 
recent studies corer et al. 1992, Dutson et al. 1992, Evans et al. 
1993). 

P. celebensis has not yet been included in the world list of threatened 
birds (Collar et al. 1994). Based on our recent investigations, at least 
one criterion for this classification would be fulfilled: population 
numbers of all three subspecies are lower than 10000 mature 
individuals. Additionally, the highly fragmented habitat is still subject 
to further destruction by continued logging, shifting agriculture and 
soil erosion. We therefore propose to include P. celebensis into the 
IUCN category ‘“‘vulnerable’’. 


Acknowledgements 


The present study was initiated by the Philippine Eagle Conservation Program 
Foundation and funded by the Austrian Science Foundation (FWF-project BIO-8889). 
The authors are very much obliged to the curators of the following museum collections 
for access to specimens in their care: The Natural History Museum (BMNH), Bird 
Department (Tring, U.K.), Universitets Zoologiske Museum (UMZC, Kobenhavn, 


A. Gamauf & M. Preleuthner 100 Bull. B.O.C. 1998 118(2) 


DK), Ryksmuseum van Natuurlijke Historie (RMNH, Leiden, NL), Zoologisches 
Museum der Humboldt Universitat Berlin (ZMB, Berlin, D), American Museum of 
Natural History (AMNH, New York, U.S.A.), Smithsonian Institution (USNM, 
Washington, DC, U.S.A.), Field Museum of Natural History (FMNH, Chicago, 
U.S.A.), Cincinnati Museum of Natural History (CMNH, Cincinnati, U.S.A.), 
Delaware Museum of Natural History (DMNH, Wilmington, U.S.A.), National 
Museum of the Philippines (PNM, Manila, PH), University of the Philippines at Los 
Banos (UPLB, Los Banos, PH), and Zoological Garden Manila (Manila, PH). We are 
especially grateful to R. Prys-Jones, P. Colston, J. Fjeldsa, C. Smeenck and R. Dekker, 
B. Stephan, G. F. Barrowclough and P. Sweet, D. Zusi, D. Willard and P. Baker, R. S. 
Kennedy and J. Brown, G. Hess, P. C. Gonzales, A. Dans and R. A. Andres for their 
helpful cooperation. We are greatly indebted to W. Pinsker, E. Bauernfeind and E. 
Eckhard for critical comments on the manuscript. 


References: 

Brooks, T. M., Evans, T. D., Dutson, G. C. L., Anderson, G. Q. A., Asane, D. C.., 
Timmins, R. J. & Toledo, A. G. 1992. The conservation status of the birds of 
Negros, Philippines. Bird Conservation International 2: 273-302. 

Brown, L. & Amadon, D. 1969. Eagles, hawks and falcons of the world. Country Life 
Books, London. 

Collar, N. J., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to watch: The world list 
of threatened birds. BirdLife International, Birdlife Conservation Series No. 4, 
Cambridge. 

Collins, N. M., Sayer, J. A. & Whitmore, T. C. 1991. The conservation atlas of tropical 
forests—Asia and the Pacific. Macmillan Press, London. 

Danielsen, F., Balete, D. S., Christensen, T. D., Heegaard, M., Jacobsen, O. F., Jensen, 
A., Lund, T. & Poulsen, M. K. 1993. Conservation of Biological Diversity in the 
Sierra Madre Mountains of Isabela and southern Cagayan Province, The Philippines. 
Department of Environment and Natural Resources (DENR), International Council 
for Bird Preservation (ICBP), Zoological Museum of Copenhagen University and 
Danish Ornithological Society (DOF), Manila. 

Delacour, J. & Mayr, E. 1946. Birds of the Philippines. The Macmillan Company, New 
York. 

del Hoyo, J., Elliot, A. & Sargatal, J. 1994. Handbook of the Birds of the World. Vol. 2. 
New World Vultures to Guineafowl. Lynx Edicions, Barcelona. 

Diamond, J. M. & Gilpin, M. E. 1983. Biogeographic umbilici and the origin of the 
Philippine avifauna. Ozkos 41: 307-321. 

Dickinson, E. C., Kennedy, R. S. & Parkes, K. C. 1991. The birds of the Philippines: an 
annotated check-list. British Ornithologists’ Union, Check-list no. 12. 

Dickinson, E. C., Kennedy, R. S., Read, D. K. & Rozendaal, F. G. 1989. Notes on the 
birds collected in the Philippines during the Steere expedition of 1887/1888. 
Nemouria 32: 1-19. 

duPont, J. E. 1971. Philippine birds. Monograph Series No. 2, Delaware Museum of 
Natural History, Greenville. 

Dutson, G. C. L., Evans, T. D., Brooks, T. M., Asane, D. C., Timmins, R. J. & Toledo, 
A. G. 1992. The conservation status of the birds of Mindoro, Philippines. Bird 
Conservation International 2: 303-325. 

Evans, T. D., Dutson, G. C. L. & Brooks, T. M. (eds) 1993. Cambridge Philippines 
rainforest project 1991: final report. Study report no. 54. Birdlife International, 
Cambridge. 

Gamauf, A., Preleuthner, M. & Winkler, H. 1998. Philippine birds of prey: interrelations 
among habitat, morphology, and behavior. Auk 115. 

Goodman, S. M. & Gonzales, P. C. 1990. The birds of Mt. Isarog National Park, 
Southern Luzon, Philippines, with particular reference to altitudinal distribution. 
Fieldiana (Zoology) 60: 1-39. 

Hauge, P., Terborgh, J., Winter, B. & Parkinson, J. 1986. Conservation priorities in the 
Philippine Archipelago. Forktail 2: 83-91. 

Heaney, L. R. 1985. Zoogeographic evidence for middle and late Pleistocene land bridges 
to the Philippine Islands. Mod. Quaternary Res. SE Asia 9: 127-143. 

Preleuthner, M. & Gamauf, A. 1998. A new subspecies of the Philippine Hawk-Eagle 
(Spizaétus philippensis) and its future prospect. 7. Raptor Res. 32. 


A. Delestrade 101 Bull. B.O.C. 1998 118(2) 


Sclater, W. L. 1919. Descriptions of new Hawks: Spizaétus nipalensis fokiensis, Spilornis 
cheela ricketti, S. c. kinabaluensis, S. c. palawanensis, Pernis celebensis steerei, with a 
synopsis of the genus Spilornis. Bull. Brit. Orn. Cl. 11: 37-42. 

Stresemann, E. 1940. Zur Kenntnis der Wespenbussarde (Pernis). Archiv fiir 
Naturgeschichte, N.F. 9: 137-193. 

Wallace, A. R. 1868. On the raptorial birds of the Malay Archipelago. bis (2) 4: 1-27. 

Weick, F. 1980. Die Greifvégel der Welt. Parey Verlag, Hamburg. 


Address: Konrad Lorenz-Institut fiir Vergleichende Verhaltensforschung, Oster- 
reichische Akademie der Wissenschaften, Savoyenstrasse 1A, A-1160 Wien, 
Austria. Present address of AG: Naturhistorisches Museum Wien, Zool. Abteilung, 
Vogelsammlung Burgring 7, A-1014 Wien, Austria. 


© British Ornithologists’ Club 1998 


Distribution and status of the Ethiopian 
population of the Chough Pyrrhocorax 
pyrrhocorax baileyi 


by Anne Delestrade 
Received 5 Fuly 1997 


The Chough Pyrrhocorax pyrrhocorax has a wide distribution, from the 
Palearctic to the Afrotropics. However, this distribution is highly 
fragmented, with numerous small isolated populations, such as in 
Scotland, Wales, the Canary Islands, several Mediterranean islands, 
Algeria and Ethiopia (sée, e€.9-} Cramp & Perrins 1994). It inhabits 
mountain areas in the main part of its range, with only the western 
populations living on sea cliffs and inshore islands. The Chough 
population in Europe is currently declining (Monaghan 1988, Cramp & 
Perrins 1994, Tucker & Heath 1994, Madge & Burn 1995), but the 
status of Chough populations outside Europe has never been accurately 
assessed. 

The Ethiopian papules is of particular interest because it is the 
southernmost (the Bale mountains are some 850km north of the 
equator), and the only population living within the Afroalpine 
ecosystem. This population is currently completely isolated from the 
others, and has been classified as a distinct subspecies, P. p. baileyi 
(Rand & Vaurie 1955). There are very limited data from Ethiopia 
(Brown 1967, Cramp & Perrins 1994), and the precise distribution and 
the size of the population are currently unknown. 


Methods 


The field study was carried out in Ethiopia between 16 November 1996 
and 18 January 1997, i.e. the dry season and harvesting time. During 
this season, Choughs are not breeding (pers. obs.), and thus gather in 
large flocks (see Blanco et al. 1993). Flocks at roosting or at foraging 


A. Delestrade 102 Bull. B.O.C. 1998 118(2) 


ERITREA 


0 300 km 
———— eee 


y Simien ey 


Mountains 


SUDAN 
DJIBOUTI 


@® Distribution of the Chough 


SOMALIA 


KENYA 
Figure 1. Distribution of the Chough in Ethiopia. 


sites were censused by visiting most suitable high-altitude massifs: Mt 
Choke (Gojam region, 4,052 m), Mt Guna (Gondar, 4,135 m), Dilanta 
highlands (Welo, 3,601 m), Mt Abune Yosef (Welo, 4,284 m), Simien 
Mountains (Gondar, Mt Ras Deshen 4,533 m), and Bale Mountains 
(Bale, Mt Tullu Deemtu 4,377 m) (Fig. 1). Estimates of population size 
were derived from observations of flock size, flock movements, and 
identification of areas used by each flock. Roosts were recorded by 
observations of flock movements on cliffs just before sunset. I spent 14 
days (7-20 December 1996) in the Simien Mountain National Park and 
20 days (27 December 1996-15 January 1997) in the Bale Mountain 
National Park in order to obtain exhaustive censuses in these two areas. 


Results 


Distribution 

During this study, the presence of Choughs was recorded in four 
difference massifs: Bale Mountains and Simien Mountains, where they 
were already known (Brown 1967, Urban & Brown 1971); The Dilanta 
highlands, from where there had been an early report by Blanford 
(1870); and a new area, the massif of Mt Abune Yosef in the Welo 
region. Choughs were not observed at Mt Choke (Gojam region) nor at 
Mt Guna (Gondar), and farmers from these areas were not familiar 
with the species, which suggests that Choughs are absent from these 
mountains. 


A. Delestrade 103 Bull. B.O.C. 1998 118(2) 


In view of the apparently suitable habitat, Choughs could be present 
in the Amba Farit Mountains (Welo region, 4,247 m, Fig. 1) but a 
search could not be arranged there. In the northern part of the Bale 
mountains, Choughs were reported by local people near Agarfa 
(Fig. 1), but their presence was not confirmed in this study. 


Flock and population size 

Three roosts in the Simien Mountain National Park were frequented 
respectively by 80, 150 and 170 individuals. One roost located in the 
Abune Yosef massif (near Lalibela, Fig. 1) was visited by 25 
individuals. Lastly, in the Bale Mountain National Park, large flocks 
gathered from different sites at dusk in one area of the Harenna 
escarpment (Rafu area), and were estimated at more than 100 
individuals, although the precise location of the roost could not be 
identified. For all areas combined, the average foraging flock size was 
60 (range 9-150, s.d. 35.5, n=26). 

Population sizes could be estimated precisely only for the Simien and 
Bale Mountains National Parks, where careful counts were made. 
Based on flock locations and sizes, the Simien Mountain National Park 
(179 km*) population was estimated at 350-500 birds, while the 
population of the Bale Mountain National Park (2, 471 km?, but with 
the Harenna forest excluded only some 1,700 km?” suitable) was 
estimated at 250-400 birds. Thus, the total population of Choughs in 
Ethiopia living within the Parks was between 600 and 900 birds. 

The minimum size of the Ethiopian Chough population based on the 
present counts would be in the order of 675-975 birds. As all areas 
outside the Simien and Bale Mountain National Parks, known to be 
inhabited by Choughs, were not checked in this survey, a total of 
1,000—1,300 birds in Ethiopia is possibly a more likely figure. One is 
obliged to wonder how these isolated populations survive at very low 
densities, and what controlling factors are involved. 


Habitat 

In Ethiopia, Choughs live in high mountains. During this study, the 
26 foraging flocks observed were in open habitats between 2,800 and 
4,200 m. Five roost sites were found in cliffs, between 3,000 and 
3,900 m. Although outside the breeding season, many pairs were 
visiting nest-sites, all in cliffs between 3,000 and 4,300 m. Foraging 
Choughs were noted in the following habitats: grazing areas, field crops 
(mainly barley and wheat), afroalpine belt, escarpments and cliffs. 
During harvesting of the cereal crops, Choughs fed on the seeds left on 
the ground. Overall therefore, Choughs depended mainly on grassland 
areas and field crops for foraging, and on cliffs for roosting and 
breeding. 


Discussion 


This study has provided distributional data and an estimate of 
population size for the Chough in Ethiopia, a first step toward the 
analysis of long-term trends in this population. Recognised as an 


A. Delestrade 104 Bull. B.O.C. 1998 118(2) 


endemic subspecies, the Ethiopian population is the most isolated one, 
the nearest other population being in southern Iran (Desfayes & Praz 
1978), about 3,000 km distant. Calls of the Ethiopian birds were found 
to differ substantially from Choughs living in the Alps (pers. obs.); a 
detailed analysis of recording will be published elsewhere. Even within 
Ethiopia, Choughs may be divided into three sub-populations, each 
composed of several hundred individuals, two in the north in the West 
Highlands (Simien and Welo Mountains), and one in the South-East 
Highlands (Bale Mountains). The two northern populations are 
separated by 200 km of unsuitable habitat, and the Bale population is 
600 km distant from the nearest northern population. Chough density 
was higher in the Simien Mountains National Park than in the Bale 
Mountains National Park (1.9-2.8 birds/km? versus 0.1-0.2 birds/km/? 
respectively), probably as a consequence of a greater availability of nest 
and roost sites in cliffs in the Simien Mountains. Obviously more data 
are needed to assess the presence of Choughs in other massifs (e.g. 
Amba Farit in Welo region). With a total population of about 1,000 
birds, and given the complete isolation from other populations, as well 
as the highly fragmented distribution within Ethiopia, it is very likely 
that exchanges of individuals between sub-populations are few (if they 
occur at all) as Choughs are known to be highly sedentary, and thus the 
status of this Ethiopian population appears to be vulnerable if not 
threatened. 


Acknowledgements 


I thank the British Ecological Society, the British Ornithologists’ Union and the Percy 
Sladen Memorial Trust, whose grants partly covered the expenses of this study. Coquoz 
Sport and Patagonia generously provided mountain equipment. I also thank the 
Ethiopian Wildlife Conservation Organisation which gave administrative support to the 
study in Ethiopia. Thanks are due also to the staffs of the Bale and Simien Mountain 
National Parks, for facilitating work in the Parks, and especially to Derbie Deksios who 
helped me in the field during all my research inside the Simien Mountain National Park. 
I lastly thank Leo Production for hospitality in their camp in Bale Mountain National 
Park, John Ash, M. Clouet, Yilma Dellelegn and Akele Yemane for information and help 
and V. Bretagnolle, D. W. Snow and an anonymous referee for their criticisms of the 
manuscript. 


References: 

Blanco, G., Fargallo, J. A. & Cuevas, J. A. 1993. Seasonal variations in numbers and 
levels of activity in a communal roost of Choughs Pyrrhocorax pyrrhocorax in central 
Spain. Avocetta 17: 41-44. 

Blanford, W. T. 1870. Observations on the Geology and Zoology of Abyssinia. Macmillan & 
Co., London. 

Brown, L. H. 1967. The occurrence of the Chough Pyrrhocorax pyrrhocorax in the 
Mendebo-Araenna Mountains of the Bale Province, Ethiopia. [bis 109: 275. 

Cramp, S. & Perrins, C. 1994. Handbook of the Birds of Europe the Middle East and North 
Africa. Vol. 8. Oxford Univ. Press. 

Desfayes, M. & Praz, J. C. 1978. Notes on habitat and distribution of montane birds in 
Southern Iran. Bonn. Zool. Beitr. 29: 18-37. 

Madge, S. & Burn, H. 1995. Crows and Jays. Christopher Helm, London. 

Monaghan, P. 1988. The background to Chough studies in Britain. In: E. Bignal & D. J. 
Curtis (eds), Choughs and Land-use in Europe. Scottish Chough Study Group, 
Argyll. i 

Rand, A. L. & Vaurie, C. 1955. Bull. Brit. Orn. Cl. 75: 28. 


R. Demey & L. D. C. Fishpool 105 Bull. B.O.C. 1998 118(2) 


Tucker, G. M. & Heath, M. F. 1994. Birds in Europe: their conservation status. BirdLife 
International (BirdLife Conservation Series no. 3), Cambridge, U.K. 

Urban, E. K. & Brown, L. H. 1971. A Checklist of the Birds of Ethiopia. Addis Ababa 
University Press, Addis Ababa. 


Address: Anne Delestrade, Centre de Recherches sur les Ecosystémes d’Altitude 
(CREA), 400 Route du Tour, Montroc, 74400 Chamonix, France. 


© British Ornithologists’ Club 1998 


On the existence of a melanistic morph of the 
Long-tailed Hawk Urotriorchis macrourus 


by Ron Demey & L. D.C. Fishpool 
Received 21 Fuly 1997 


The Long-tailed Hawk Urotriorchis macrourus (Hartlaub, 1855) is a 
little recorded raptor occurring in the Upper and Lower Guinea forest 
blocks of tropical Africa. New information concerning its distribution 
has come to light since the publication of The Birds of Africa (Brown 
et al. 1982) and its range is now known to extend from eastern 
Sierra Leone (Gola forest), Liberia and southeastern Guinea (Ziama 
forest) through Ivory Coast and Ghana to western ‘Togo, and from 
western Nigeria and Cameroon southwards to Equatorial Guinea, 
Gabon, Congo, Cabinda and former Zaire, and eastwards to 
southwestern Central African Republic, just reaching the extreme 
south of Sudan and Bwamba in western Uganda (Allport et al. 1989, 
Britton 1980, Brown et al. 1982, Carroll 1988, Cheke & Walsh 1996, 
Dowsett-Lemaire & Dowsett 1989, Elgood et al. 1994, Halleux 1994, 
Hillman & Hillman 1986). 

The species is monotypic; the formerly recognised race batesz, 
supposed to have a longer tail, and proposed for populations from 
Cameroon eastwards (Bates 1930, Dekeyser & Derivot 1966, 
Mackworth-Praed & Grant 1970) is now considered invalid (Brown 
et al. 1982, Kemp 1994). 

The purpose of this note is to draw attention to some conflict or 
inconsistency that exists in the literature concerning the colouration of 
this species, which emphasises how poorly known it is. 

The general colouration of the adult is entirely dark grey on the 
upperparts with contrasting white uppertail-coverts; the exceptionally 
long, graduated tail is black tipped and barred white. Below, it is paler 
grey on the throat, with the remainder of the underparts and the 
underwing-coverts, typically, a rich chestnut brown; undertail-coverts 
are white. This much is non-controversial. There is, however, dispute 
over the existence of the so-called melanistic morph of this species in 
which the chestnut is replaced by a dark slaty grey. 

The melanistic morph was first described by Sharpe (1870) from a 
specimen, sex unspecified, secured for the Norwich Museum, which 


R. Demey & L. D. C. Fishpool 106 Bull. B.O.C. 1998 118(2) 


apparently constituted the first example of this species to have reached 
Britain, the type (a juvenile) having been sent to the Leiden Museum 
by Pel, provided by ‘Temminck and described by Hartlaub in 1855. An 
accurate colour lithograph by Keulemans illustrates Sharpe’s paper. 
For reasons now unclear, four years later Sharpe (1874) considered the 
colour difference to be sex related, with the grey morph being male, the 
chestnut form female. This was repeated by Reichenow (1901). 
Bannerman (1930) mentions having examined the melanistic specimen 
from the Norwich Museum and gives a description of it (“breast and 
belly entirely grey, a faint indication of reddish-brown feathers 
appearing on the throat and here and there on the breast’’). He does not 
mention, however, the existence of this morph in the shorter, 
two-volume version of his work (Bannerman 1953). It is not mentioned 
either by Bates (1930), Lippens & Wille (1976), Mackworth-Praed & 
Grant (1952), Serle, Morel & Hartwig (1977), Snow (1978) and 
Williams & Arlott (1980). In the forty years between Bannerman (1930) 
and Mackworth-Praed & Grant (1970), Schouteden (1954) seems to be 
the only author to make explicit reference to it, stating that “‘the breast 
is sometimes grey’’. Not to mention it is, of course, not to deny its 
occurrence—one might infer that in some cases lack of space or 
superficial treatment may have precluded mention. This seems, 
however, to have had the unfortunate result of leading others at least to 
overlook or ignore its existence. 

Thus, Brown et al. (1982) describe the underparts of the adult as 
being wholly chestnut, and do not mention the melanistic morph. This 
is surprising, however, because Mackworth-Praed & Grant (1970) 
clearly state that the underparts may be either chestnut or blackish slate 
and include a colour illustration of both morphs. It is even more 
remarkable in view of the fact that the principal author of the former, in 
another major work (Brown & Amadon 1968), illustrates both colour 
morphs in a plate but, curiously, omits any mention of the melanistic 
phase in the text. Recently Kemp (1994), although mentioning that “‘a 
melanistic morph has been claimed to exist and even been depicted’, 
has gone so far as to conclude that this was “apparently 
unsubstantiated’. 

The Natural History Museum (Tring) holds 23 adult specimens of 
which two are grey morphs. One specimen (reg. no. 1955.6.N20.3245) 
was that originally held in the Norwich Museum and illustrated in 
Sharpe’s paper (it bears a label to this effect). Locality data are given 
simply as Denkera, Fantee County [Ghana]. The second specimen 
(reg. no. 1938.4.6.3) was collected on 18 January 1938 at Ondo, 
Nigeria. No other data are given. Among the 21 adult specimens in the 
Royal Museum for Central Africa (Tervuren, Belgium), all except one 
of which are from the Lower Guinea forest block in former Zaire, no 
grey morphs are present. Neither of the grey specimens in Tring is 
sexed, making Sharpe’s (1874) claim for the dimorphism being 
sex-linked the more intriguing. Brown et al. (1982) state that females 
average larger than males, although the measurements they give are for 
both sexes combined. The range for wing length is stated to be 
266-310 mm (sample size unspecified). Wing lengths of the grey 


R. Demey © L. D.C. Fishpool 107 Bull. B.O.C. 1998 118(2) 


individuals from Ghana and Nigeria measured 305 and 268 mm 
respectively. While not conclusive this might be taken to suggest that 
the grey morph occurs in both sexes. 

In Ivory Coast, the Long-tailed Hawk is not uncommon in Yapo 
Forest, 5°42'N 4°6’W (Demey & Fishpool 1994). Although we regularly 
heard the species there, we rarely saw it. Indeed, during 167 visits to 
Yapo forest over the course of five years we observed the Long-tailed 
Hawk on 18 occasions only, involving 22 individuals: two of these 
were seen to be melanistic birds, although the proportion could have 
been slightly higher since in some cases (number unrecorded) the 
observations concerned rear views of birds in flight. Brief details of 
these sightings are as follows. On 25 May 1986, RD observed a bird 
corresponding to the description of the melanistic morph (Mackworth- 
Praed & Grant 1970). The bird crossed a clearing at less than 10m 
from the observer, and landed in a tree. It was not shy, allowing close 
and detailed observation through a telescope of 20 X magnification 
during more than 15 minutes. Notes taken on the spot describe the 
underparts as slate-grey, concolorous with the upperparts; upper- and 
undertail-coverts pure white; tail very long, black with white spots, 
looking somewhat worn; underwing barred. Eye yellow; bill 
greyish-black; legs yellow. A second melanistic individual was seen 
under similar circumstances at a different locality in Yapo by LDCF on 
14 February 1988. The plumage description taken at the time matches 
the foregoing exactly. The only addition was that the cere of this 
individual was noted as grey. 

Another record of this morph has been documented from the Lower 
Guinea forest block, in Gabon. Brosset & Erard (1986) report a 
sighting, in April 1985, of a ‘mixed’ pair, of which one of the birds 
showed uniformly grey underparts. 

It thus appears that the claims of the existence of a melanistic morph 
of the Long-tailed Hawk are indeed well founded. 


Acknowledgements 


We are grateful to Dr Robert Prys-Jones and Michael Walters at the Natural History 
Museum (Tring) and Dr Michel Louette at the Royal Museum for Central Africa 
(Tervuren) for access to skins. Dr Louette is also thanked for his help in the search for 
relevant literature. 


References: 

Allport, G., Ausden, M., Hayman, P. V., Robertson, P. & Wood, P. 1989. The 
Conservation of the Birds of Gola Forest, Sierra Leone. Study Report 38, International 
Council for Bird Preservation, Cambridge. 

Bannerman, D. A. 1930. The Birds of Tropical West Africa. Vol.1. Crown Agents, 
London. 

Bannerman, D. A. 1953. The Birds of West and Equatorial Africa. Vol. 1. Oliver & Boyd, 
Edinburgh. 

Bates, G. L. 1930. Handbook of the Birds of West Africa. Bale & Danielsson, London. 

Britton, P. L. (ed.) 1980. Birds of East Africa. EANHS, Nairobi. 

Brosset, A. & Erard, C. 1986. Les Oiseaux des Régions Forestiéres du Nord-Est du Gabon, 
vol. 1: Ecologie et Comportement des Espéces. Société Nationale de Protection de la 
Nature, Paris. 

Brown, L. & Amadon, D. 1986. Eagles, Hawks and Falcons of the World. Vol. 2. Hamlyn, 
Middlesex. 


CT) Collins 108 Bull. B.O.C. 1998 118(2) 


Brown, L. H., Urban, E. K. & Newman, K. 1982. The Birds of Africa. Vol. 1. Academic 
Press, London. 

Carroll, R. W. 1988. Birds of the Central African Republic. Malimbus 10: 177-200. 

Cheke, R. A. & Walsh, J. F. 1996. The Birds of Togo. B.O.U. Check-list no. 14. British 
Ornithologists’ Union, ‘Tring. 

Dekeyser, P. L. & Derivot, J. H. 1966. Les Oiseaux de V’OQuest Africain. Fasc. I. 
Initiations et Etudes africaines No. XIX. Inst. fon. Afr. noire, Dakar. 

Demey, R. & Fishpool, L. D. C. 1994. The birds of Yapo Forest, Ivory Coast. Malimbus 
16: 100-122. 

Dowsett-Lemaire, F. & Dowsett, R. J. 1989. Liste commentée des oiseaux de la forét du 
Mayombe (Congo). Pp. 5—16 im R. J. Dowsett & F. Dowsett-Lemaire (eds), Enquéte 
faunistique dans la forét du Mayombe et check-liste des oiseaux et des mammiféres 
du Congo. Tauraco Res. Rep. 2. 

Elgood, J. H., Heigham, J. B., Moore, A. M., Nason, A. M., Sharland, R. E. & Skinner, 
N. J. 1994. The Birds of Nigeria. B.O.U. Check-list no.4 (2nd edn). British 
Ornithologists’ Union, Tring. 

Halleux, D. 1994. Annotated bird list of Macenta Prefecture, Guinea. Malimbus 16: 
10-29. 

Hillman, J. C. & Hillman, S. M. 1986. Notes on some unusual birds of the Bangangai 
area, South-west Sudan. Scopus 10: 29-31. 

Kemp, A. C. 1994. Long-tailed Hawk Urotriorchis macrourus. P. 164 in J. Del Hoyo, A. 
Elliott & J. Sargatal (eds), Handbook of the Birds of the World. Vol. 2. Lynx Edicions, 
Barcelona. 

Lippens, L. & Wille, H. 1976. Les Oiseaux du Zaire. Lannoo, Tielt. 

Mackworth-Praed, C. W. & Grant, C. H. B. 1952. Birds of Eastern and North Eastern 
Africa. Vol. 1. Longman, London. 

Mackworth-Praed, C. W. & Grant. C. H. B. 1970. Birds of West Central and Western 
Africa. Vol. 1. Longman, London. 

Reichenow, A. 1901. Die Vogel Afrikas. Vol. 1. Verlag von J. Neumann, Neudamm. 

Schouteden, H. 1954. Faune du Congo Belge et du Ruanda-Urundi. III. Oiseaux Non 
Passereaux. Ann. Mus. Congo belge, 8°, Sc. Zool., vol. 29. 

Serle, W., Morel, G. J. & Hartwig, W. 1977. A Field Guide to the Birds of West Africa. 
Collins, London. 

Sharpe, R. B. 1870. On a fourth collection of birds from the Fantee country. bis (2) 6: 
52-59. 

Sharpe, R. B. 1874. Catalogue of the birds in the British Museum. Vol. 1. British Museum 
(Natural History), London. 

Snow, D. W. (ed.) 1978. An Atlas of Speciation in African Non-Passerine Birds. British 
Museum (Natural History), London. 

Williams, J. G. & Arlott, N. 1980. A Field Guide to the Birds of East Africa. Collins, 
London. 


Addresses: Ron Demey, Grote Peperstraat 3, B-9100 Sint-Niklaas, Belgium. Dr L. D. C. 
Fishpool, BirdLife International, Wellbrook Court, Girton Road, Cambridge 
CB3 ONA, U.K. 


© British Ornithologists’ Club 1998 


Food delivery and chick provisioning in 
cypseloidine swifts 


by Charles T. Collins 
Received 9 August 1997 


Swifts (Apodidae) catch all of their arthropod food on the wing. These 
items, mostly insects, are brought back to nestlings in two strikingly 


C. T. Collins 109 Bull. B.O.C. 1998 118(2) 


different ways which in turn affect the kind of food adults capture, their 
foraging range and chick provisioning rate. 

Swifts and swiftlets in the subfamilies Chaeturinae and Apodinae 
bring the food items back to nestlings in a consolidated mass or 
bolus glued together with saliva and carried in the mouth. For Apus 
apus, an individual bolus typically weighs 0.70-1.75 g, occasionally 
2.0—2.5 g (Lack & Owen 1955, Martins & Wright 1993), and contains 
90-850 individual prey items (Lack & Owen 1955, Collins 
unpublished). A bolus can contain as many as 1500 very small 
insects such as aphids (Homoptera, Aphididae) (Lack & Owen 1955). 
Although food boluses can occasionally consist of only one or a few 
types of insects, they typically contain representatives of numerous 
orders and families of insects and ballooning spiders (Hespenheide 
1975, Collins 1968, Tarburton 1986a, Bull & Beckwith 1993). The 
food bolus causes a visible distension of the floor of the mouth which 
is observable in swifts returning to feed nestlings (Lack 1956, 
Arn-Willi 1960, Cramp 1985). The adult inserts its bill into the open 
mouth of the begging chick (Lack 1956, Arn-Willi 1960) and usually 
passes most or all of the bolus to a single older chick; the bolus may 
be divided among several very small chicks (Lack & Lack 1951). 
Swifts are efficient foragers and can rapidly gather a bolus of food. 
Lack & Lack (1951) report individual Apus apus gathering 1.2 g and 
1.7 g of insects in 47 and 64 minutes respectively. Feeding rates in 
the Chaeturinae and Apodinae are highly variable, ranging from 3-4 
feedings per hour to 4.6-10.7 feedings per 10 hours (Moreau 
1942a,b, Lack & Lack 1951, Collins 1968, Tarburton 1986b, Bull & 
Collins 1993). Feeding rates are correlated with both brood size and 
weather-related differences in food abundance (Lack & Lack 1951, 
Lack & Owen 1955, Tarburton 1986b). The mode of food delivery, 
boluses carried in the mouth, directly limits the amount of food 
which can be brought back per feeding trip and, indirectly, the 
effective foraging range. It would be inefficient for such birds to 
expand their foraging range to the extent that excessive time and 
energy are spent in transport of individual boluses from distant 
foraging areas. The Ecological Cost of Transportation (Garland 
1983) or “percent of total daily energy expenditure which is 
consumed by locomotion’? to and from foraging areas (Whitacre 
1992) would become unacceptably high. Additional theoretical 
discussion of foraging efficiency in birds and the trade-offs between 
load size and foraging distance is presented by Krebs & Davies 
(1993). 

The 12-13 species of New World swifts in the genera Cypseloides and 
Streptoprocne (Sibley & Monroe 1990, Chantler & Driessens 1995) have 
many distinctive morphological and breeding characteristics warrant- 
ing their separation as the subfamily Cypseloidinae (Brooke 1970). 
Included is their tendency to nest in dark damp locations, frequently 
near or behind waterfalls (Knorr 1961, Snow 1962, Collins 1968, Marin 
& Styles 1992). It has also been noted that the rate of food delivery to 
chicks is notably low. In the Chestnut-collared Swift Cypseloides 
rutilus, feeding intervals were usually longer than 100 minutes (Collins 


C. T. Collins 110 Bull. B.O.C. 1998 118(2) 


1968). Similarly, in the Black Swift Cypseloides niger Michael (1927) 
noted several hours between feedings and possibly only a single feeding 
late in the day or at dusk. The larger White-naped Swift Streptoprocne 
semicollaris and White-collared Swift Streptoprocne zonaris also appear 
to make only a single feeding trip to the nest per day (Whitacre 
1992); 

Another subfamiulial difference which has not been given attention is 
the mode of food delivery to chicks. The cypseloidine swifts do not 
carry food for the chicks as a saliva-coated bolus in the mouth, but as an 
unconsolidated mass in the oesophagus. The distended oesophagus in 
White-naped and White-collared Swifts was 100-135 mm long, 26 mm 
in diameter, and contained 538-1078 insects 80.7% of which were flying 
ants (Formicidae: Azteca, Solenopsis) (Rowley & Orr 1962, 1965). 
‘There also appeared to be a “fringed valve situated immediately behind 
the glottis’ (Rowley & Orr 1962). The mean weight of the oesophagus 
contents of White-collared and White-naped Swifts was 5.0 g and 6.7 g, 
and they contained up to 1044 and 1218 prey items respectively 
(Whitacre 1992). Collins & Landy (1968 reported masses of 
Hymenoptera (72.4% of one species of winged ant) in the “‘throat’’ 
(=oesophagus) of two adult Black Swifts collected at night near nests in 
Veracruz, Mexico. Captured individuals of several swifts in both 
Cypseloides and Streptoprocne readily regurgitated masses of insect 
food, 35-100% again being winged ants (Foerster 1987, Marin & Styles 
1992, Whitacre 1992, Collins unpublished). Other swarming insects as 
fig-wasps (Hymenoptera, Blastophagidae) and termites (Isoptera) were 
also prominent in some food samples (Collins 1968, Whitacre 1992, 
Marin & Styles 1992). 

The preponderance of swarming insects, particularly lipid-rich 
winged ants, in the diets of Cypseloides and Streptoprocne swifts ranging 
in size from 20 to 180 g body mass suggests that the Cypseloidinae are 
foraging specialists feeding on prey that occur in dense, but possibly 
widely dispersed, patches. These swifts would be expected to forage 
over wider ranges than other swifts (Chaeturinae, Apodinae) to locate 
such food sources. This seems to be true for several species of 
Cypseloides (Collins pers. obs.) and has been confirmed by radio 
telemetry for Streptoprocne (Whitacre 1992). The ability to bring back 
in the oesophagus larger quantities of food, than could be 
accommodated as individual boluses carried in the mouth, would seem 
to involve a critical morphological adaptation enabling these swifts to 
utilize both distant and patchily distributed prey resources. I suggest 
that a single large mass of food carried in the oesophagus can be used to 
provision a chick in the form of multiple feedings at one time, or 
sequential feedings over an extended period of time, thereby 
compensating for the infrequent feeding trips to the nest which is 
typical of these swifts. 

Recent observations support these suggestions. On 19 July 1997, at 
a breeding colony of Black Swifts located at Mosse Brae Falls 
near Dunsmuir, Siskiyou Co., California, an adult returned to the 
nest at about 19.40 local time. Shortly afterwards it was seen to 
make open-mouth gaping and stretching motions. The mouth was 


C. T. Collins 111 Bull. B.O.C. 1998 118(2) 


clearly empty of food at this time. Following this, the floor of the 
then closed mouth bulged out with regurgitated food which was 
promptly fed to a half-grown nestling. In rapid succession, over the 
next 2-3 minutes 10 more similar sized regurgitations were fed to the 
chick. In each case, prior to feeding the chick, the floor of the mouth 
appeared distended, approximately to the same extent as in swifts 
carrying a single food bolus, and empty afterwards. In total, the chick 
appeared to receive, in one bout of provisioning, the equivalent mass 
of food as many individual boluses delivered throughout the day. 
Multiple feedings over a period of eight minutes has also been 
observed in British Colombia (Grant 1966). On 8 August 1987, at a 
nesting site at Lawler Falls in southern California (Foerster & Collins 
oy. -Cellins & _ Foerster 1995) -an adult. Black Swift’ ‘was 
photographed feeding its chick at dusk shortly after returning to the 
nest for the night. What appeared to be the same adult again fed the 
chick over an hour later, well after dark (Collins & Peterson 1998). 
Thus multiple feedings over a longer period of time are also 
confirmed. Elsewhere, Black Swifts collected after dark, at a nest, had 
large quantities of food in their oesophagus (Collins & Landy 1968). 
This also suggests that the chick would be provisioned one or more 
times during the night. 

Hespendeide (1975) was perhaps the first to suggest that some swifts 
might be specialists on a limited array of species, particularly 
calorically-dense swarming insects. However, he suggested that this 
was limited to a few of the very large species, as those in Streptoprocne. 
Whitacre (1992) verified that swarm feeding was typical of both 
White-collared and White-naped Swifts but related it mostly to 
patchily distributed food resources in seasonally dry climates and the 
development of coloniality. I think there is now enough information on 
swarm feeding by all species of the Cypseloidinae studied to date to 
suggest a stronger phylogenetic component to this behavioural 
specialization, and its morphological correlate, of carrying larger 
quantities of food in the expanded oesophagus and provisioning chicks 
at longer intervals. Swarm feeding, however, may not be limited to the 
Cypseloidinae but also appears in several species of swiftlets 
(Aerodramus) (Harrison 1976, Collins & Francis, unpublished). Studies 
of the foraging range and feeding rate in these species are clearly called 
for. 


References: 

Arn-Willi, H. 1960. Biologische Studien am Alpensegler. Verlag Vogt-Schild, Solothurn. 

Brooke, R. K. 1970. Taxonomic and evolutionary notes on the subfamilies, tribes, 
genera and subgenera of the swifts (Aves: Apodidae). Durban Mus. Novit. 9: 
13-24. 

Bull, E. L. & Beckwith, R. C. 1993. Diet and foraging behavior of Vaux’s Swifts in 
northeastern Oregon. Condor 96: 1017-1923. 

Bull, E. L. & Collins, C. T. 1993. Vaux’s Swift (Chaetura vauxti). In: The Birds of North 
America, No. 77 (A. Poole & F. Gill, eds). Academy of Nat. Sci., Philadelphia. 
Chantler, P. & Driessens, G. 1995. A Guide to the Swifts and Treeswifts of the World. Pica 

Press, Sussex, U.K. 
Collins, C. T. 1968. The comparative biology of two species of swifts in Trinidad, West 
Indies. Bull. Fla. St. Mus. 11;: 257-320. 


Cyan! Catlins Lt? Bull. B.O.C. 1998 118(2) 


Collins, C. 'T. & Foerster, K. S. 1995. Nest site fidelity and adult longevity in the Black 
Swift (Cypseloides niger). North American Bird Bander 20: 11-14. 

Collins, C. T. & Landy, M. J. 1968. Breeding of the Black Swift in Veracruz, Mexico. 
Bull. So. Calif. Acad. Sci. 67: 266-268. 

Collins, C. T. & Peterson, B. M. 1998. Nocturnal chick provisioning in Black Swifts. 
West. Birds, in press. 

Cramp, W. (ed.). 1985. The Birds of the Western Palearctic. Vol. 4. Oxford University 
Press. 

Foerster, K. S. 1987. The distribution and breeding biology of the Black Swift 
(Cypseloides niger) in southern California. M.S. Thesis, California State University, 
Long Beach. 

Foerster, K. S. & Collins, C. T. 1990. Breeding distribution of the Black Swift in 
southern California. West. Birds 21: 1-9. 

Garland, T., Jr. 1983. Scaling the ecological cost of transport to body mass in terrestrial 
mammals. Am. Nat. 121: 571-587. 

Grant, J. 1966. A Black Swift nest in British Colombia. Canad. Field-Nat. 80: 60-61. 

Harrisson, 'T’. 1976. Food of Collocalia swiftlets (Aves: Apodidae) at Niah Great Cave in 
Borneo. 7. Bombay Nat. Hist. Soc. 71: 376-393. 

Hespenheide, H. A. 1975. Selective predation by two swifts and a swallow in Central 
America. Ibis 117: 82-99. 

Knorr, O. 1961. The ecological and geographical distribution of the Black Swift in 
Colorado. Wilson Bull. 73: 155-170. 

Krebs, J. R. & Davies, N. B. 1993. An Introduction to Behavioural Ecology. 3rd edn. 
Blackwell Scientific Publications, Oxford. 

Lack, D. 1956. Swifts in a Tower. Methuen and Co., London. 

Lack, D. & Lack, E. 1951. The breeding biology of the Swift Apus apus. Ibis 93: 501-546. 

Lack, D. & Owen, D. F. 1955. The food of the Swift. ¥. Animal Ecol. 24: 120-136. 

Marin, A. M. & Styles, F. G. 1992. On the biology of five species of swifts (Apodidae, 

Cypseloidinae) in Costa Rica. Proc. West. Found. Vert. Zool. 4: 286-351. 

Martins, T. L. F. & Wright, J. 1993. Cost of reproduction and allocation of food between 

parent and young in the Swift (Apus apus). Behav. Ecol. 4: 213-223. 

Michael, C. W. 1927. Black Swifts nesting in Yosemite National Park. Condor 29: 89-97. 

Moreau, R. E. 1942a. The breeding biology of Micropus caffer streubelit Hartlaub, the 
White-rumped Swift. [bis 74: 27-49. 

Moreau, R. E. 1942b. Colletoptera affinis at the nest. Ostrich 13: 137-147. 

Rowley, J. S. & Orr, R. T. 1962. The nesting of the White-naped Swift. Condor 64: 
361-367. 

Rowley, J. S. & Orr, R. T. 1965. Nesting and feeding habits of the White-collared Swift. 
Condor 67: 449-456. 

Sibley, C. G. & Monroe, B. L., Jr. 1990. Distribution and Taxonomy of Birds of the World. 
Yale University Press. 

Snow, D. W. 1962. Notes on the biology of some Trinidad swifts. Zoologica 47: 129-139. 

Tarburton, M. K. 1986a. The food of the White-rumped Swiftlet (Aerodramaus 
spodiopygius in Fiji. Notornis 33: 1-16. 

Tarburton, M. K. 1986b. Breeding of the White-rumped Swiftlet in Fyi. Emu 86: 
214-227. 

Whiteacre, D. F. 1992. Studies of the ecology of the White-collared Swift (Streptoprocne 
zonaris) and White-naped Swift (GS. semicollaris), and of patterns of adaptation 
among the swifts (Aves: Apodidae). Unpublished Ph.D. dissertation, University 
Calif., Davis. 


Address: Dr C. T. Collins, Department of Biological Sciences, California State 
University, Long Beach, California 90840, U.S.A. 


© British Ornithologists’ Club 1998 


JF. Riley et al. 113 Bull. B.O.C. 1998 118(2) 


The taxonomic status of Halcyon enigma on 
the ‘T‘alaud islands, Indonesia 


by Fon Riley, Damien Hicks & Fames C. Wardill 


Received 21 August 1997 


The kingfisher Halcyon enigma Hartert, 1904 is endemic to the 
Talaud islands, a small archipelago located between Sulawesi and the 
Philippines. The unresolved taxonomic position of Halcyon king- 
fishers on these islands partly reflects a lack of recent fieldwork, but 
new information gathered by the authors in September and October 
1995 and January to March 1997, suggests that enigma should be 
regarded as a species, and not a subspecies of Halcyon chloris. 

On the three largest islands in the Talaud group, Karakelang, 
Salibabu and Kabaruan, kingfishers resembling Collared Kingfisher 
H. chloris have been collected, specimens of which can be fitted into 
two exclusive size ranges. The small specimens were initially thought to 
be forms of Sacred Kingfisher Halcyon sancta (Meyer & Wiglesworth 
1895) or immature H. chloris (Meyer & Wiglesworth 1898). 

These conclusions were shown to be erroneous, and the small form was 
first described as a separate species, Halcyon enigma, by Hartert (1904). 
His conclusions were based on eight specimens with bills 35-40 mm long 
against 45-50 mm in chloris specimens from the Talaud group, and wings 
94—98 mm as opposed to 108-120 mm in chloris. Hartert noted ‘“Whether 
this small form on Talaut is a geographical representative of chloris 
(though both are found on Talaut, one might only breed there, the other 
be an occasional immigrant), or a perfectly developed species coexisting 
with typical large chloris, or a local aberration—for it is only known on 
‘Talaut—it will be desirable to have a name for it’’. 

Oberholser (1919) treated both large and small forms as Halcyon 
chloris enigma, concluding that an unusual variation in size occurs, 
although he examined only one specimen of enigma. Such a large size 
dimorphism within a population of birds is unknown (Eck 1978), and 
Oberholser’s conclusion can be discounted. 

Eck (1978) reviewed the taxonomic position of the two forms, having 
for examination 12 small specimens from Karakelang and 21 large 
specimens from Karakelang, Salibabu and Kabaruan. The small birds 
were shown to be close to H. c. chloris of Sulawesi in colouration, whilst 
the larger birds were closer to H. c. collaris of the Philippines or H. c. 
teraoki of Palau. Measurements were taken from the specimens and it 
was shown that an uninterrupted series can be made from skins. Eck 
therefore concluded that the two forms on Talaud were conspecific; 
enigma represented a small form of chloris on 'Talaud, whilst larger 
birds were named Halcyon chloris ssp. 

Greenway (1978) suggested that large birds could be migrants from 
Sulawesi that had lingered on. Talaud, but the seasonal movements of 
Sulawesi and Philippine birds need further investigation (White & 
Bruce 1986). 


F. Riley et al. 114 Bull. B.O.C. 1998 118(2) 


Other reviews noted that if both forms were shown to breed on the 
islands and behavioural differences are observed, enigma would 
represent a separate species (Fry 1980, Eck 1978). Bruce visited 
Salibabu in 1978 and ‘“‘found apparent ecological separation, with one 
form in the forested areas (presumably enigma) and others in more 
characteristic coastal habitat’? (White & Bruce 1986). 

Recent authorities (Andrew 1992, Sibley & Monroe 1990) list enigma 
as a separate species. In recent years researchers have failed to record 
the species on Karakelang (Taylor 1991, Bishop 1992, D. A. Holmes in 
litt.), although Rozendaal collected 6 specimens in the period 12-25 
February 1985 (R. W. R. J. Dekker in Iitt.). | 

In 1995 and 1997 chloris-type birds were observed on both Salibabu 
and Karakelang, and our fieldwork supported the suggestion that the 
small form on 'Talaud should be regarded as a separate species, Halcyon 
enigma and the larger form as Halcyon chloris. This conclusion is based 
on new information gathered in four areas and a review of the 
published data. In the discussion that follows, small birds are named 
enigma and large birds chloris. 


Field characters 

The two forms were easily identifiable in the field by the differences 
in colouration and size noted by Eck (1978). The following description 
is based on features noted in the field. 

Emgma. Eye dark; legs dark; bill - upper mandible black, lower 
mandible basal half horn, distal half black. Clean white underparts, 
slightly washed with buff on the throat. The white extends round the 
neck to form a broad, well marked collar, bordered above by black, the 
black not extending onto ear-coverts. Small white spot on the nape. 
Crown, forehead, ear-coverts and upper nape are uniform deep 
bottle-green with a blue tinge. Loral patch is white and extends to 
reach the eye. Back and mantle are dirty olive green, contrasting with 
the green—blue of the wings. The wings are darkest on flight feathers; 
scapulars same colour as back and mantle; coverts light blue and 
slightly iridescent. Rump electric blue, lighter than iridescent blue tail. 

Chloris. In contrast has a black band extending from the eye to the 
black collar. The upperparts and wings of chloris are a uniform blue 
without green hints, and chloris has only a small white loral patch, 
which never reaches the eye. 

In the field enigma is most easily separated by its shorter bill that 
appears less heavy, partly because the lower mandible is less protruding 
and more extensively coloured horn from the base. Enigma also has a 
noticeably shorter tail compared to chloris. Enigma characteristically 
adopts a slightly hunched posture when perched, leading to a dumpy 
appearance. 


Ecological separation . 

As suggested by White & Bruce (1986), the two forms were found to 
show clear ecological separation. Enigma was found only in forest or 
forest edge habitats, and is commonest in undisturbed forest on 
Karakelang. It was also commonly encountered in degraded or 


F. Riley et al. 115 Bull. B.O.C. 1998 118(2) 


secondary forest, and so seems able to withstand some _ habitat 
alteration, although in cultivated areas it is apparently out-competed by 
chloris. 

Enigma was most commonly observed in the mid-canopy, between 6 
and 15m above ground, and was not seen in the sub-canopy, being 
replaced in this zone by the Ruddy Kingfisher Halcyon coromanda. At 
rest birds habitually perched in the mid-canopy layer, but were 
observed in the crowns of trees of heights up to 20 m. Birds were seen 
feeding along rivers and streams, as well as within forest, utilising a 
suitable vantage point to scan the ground below, before diving down 
onto prey. Prey items appeared to include small grasshoppers and river 
snails. 

Chloris was noted on Salibabu and Karakelang in coastal habitats 
typical of the species, including mangroves, cultivated areas, such as 
coconut plantations and low-lying secondary scrub habitats. The 
species was commonly observed in these areas and was even found in 
towns, perching on electricity cables. 


Sympatric breeding 

It has been suggested that chloris are migrants to Talaud from 
Sulawesi or the Philippines and enigma are resident breeders on the 
islands (Eck 1978, Greenway 1978). As noted by Fry (1980), if both 
forms are shown to be resident and breeding on ‘Talaud, enigma must 
be accorded specific status. 

There are now dated specimens and field observations of enigma 
between January and November and of chloris between January and 
November; the absence of records for either form in December simply 
reflects the fact that no ornithologists have visited Talaud in this 
month. If emigration were taking place, some seasonal variation in 
relative encounter rates would be expected; in 1995 and 1997 enigma 
and chloris were observed at similar frequencies in both years. Hence it 
may be concluded that both enigma and chloris are resident on the 
islands. 

Previous observations of chloris indicate that its breeding season is 
August—October in Sulawesi (White & Bruce 1986) and April-June in 
the Philippines (Dickinson et al. 1991). All published sightings of 
chloris on ‘Yalaud therefore fall within the known breeding seasons of 
adjacent populations, suggesting that the population is resident on 
Talaud and breeds. Evidence of a breeding population of chloris on 
Karakelang was provided when a single juvenile bird was observed near 
the village of Bengel (4°14'N, 126°49’E) on 23 September 1995. The 
bird had been captured by local children, who reported it had recently 
fledged from a site near the village. The bird was identified tentatively 
as chloris on size, in particular the large bill which, although not 
measured, even in a young bird appeared proportionately too large for 
enigma, and as a juvenile by a faint buff wash to the underparts and 
brown-buff freckling on collar and breast. The September date falls 
within the range given by White & Bruce (1986) for breeding on 
Sulawesi. In late March 1997, on a tiny offshore islet near Dapalan, 
Karakelang (4°24’N, 126°55’ E), a pair of chloris were seen displaying, 


F. Riley et al. 116 Bull. B.O.C. 1998 118(2) 


both birds perching close together on a tree branch, simultaneously 
uttering a quiet, harsh toned kur kur kur call whilst twisting heads 
from side to side, alternately stretching and pointing the bill up or 
down. 

No conclusive proof of breeding by enigma was obtained, but during 
late September and October 1995, birds were paired and holding 
territory 1n central Karakelang. Males were seen aggressively defending 
territories from encroaching individuals and on one occasion pair- 
bonding behaviour was observed; a presumed male caught a prey item, 
flew to join the female and presented the prey to her before both birds 
flew away. 


Vocalisations 

The two forms on ‘Talaud are separable by call. Emgma most 
commonly gives a repetitive kee kee kee kee kee, each note usually 
repeated five times followed by a brief pause followed by 5 more notes; 
occasionally birds give 4 or 6 repeats; there is no variation in pitch and 
all notes are of equal length. Chloris also gives a version of this call, but 
there are always more repeats, sometimes up to 20, but more usually 7 
to 10 notes and the call note is slightly disyllabic. The call of enigma is 
noticeably higher in pitch than chloris, and the note is monosyllabic. 

A second call given by enigma consists of a disyllabic kz-kac or ki-kee, 
the first note hard and grating, rising in pitch, whilst the second note 
descends in pitch slightly, is more drawn out, but cuts off sharply. This 
call is similar to a double note call commonly given by chloris, but the 
latter species always produces a stronger, harsher note of lower pitch 
and the sequence is given at a greater speed and 1s not as stuttering as 
that of enigma usually is. This characteristic double note of chloris can 
be transcribed as kee-ka, the emphasis being on the second note. 

On 2 March 1997 a pair of enigma were repeatedly chased and 
harried by an adult Black-naped Oriole Oviolus chinensis in the lower 
canopy of a 15 m tall tree. The kingfishers responded with a rapid, loud 
and monosyllabic alarm call kz ki ki ki ki ki lasting for approximately 5 
or 10 seconds. 


Discussion 

Based on the evidence from recent fieldwork, in addition to that 
already available from specimens, Halcyon enigma should be accorded 
specific status. The two forms found on Talaud are morphologically 
similar; vocalisations, plumage and size differences alone might not 
provide sufficient basis on which to separate them. However, since 
there is clear ecological separation of the forms, with both almost 
certainly breeding residents on the islands, it seems that they exist 
sympatrically and must be regarded as distinct species. 

Enigma is the only species endemic to the Talaud islands and listed as 
Near-threatened (White & Bruce 1986, Collar et al. 1994). Its 
conservation status is not of immediate concern, but recent 
developments on Karakelang, notably the establishment of a logging 
concession in the north of the island, require the species’ status to be 
monitored. Enigma is dependent on forest, but on Karakelang large 


JF. Riley et al. 117 Bull. B.O.C. 1998 118(2) 


areas of habitat still exist, with approximately 36000 ha having 
protected status (PPA 1980). In addition the species does seem to be 
able to survive in heavily degraded forest habitat. Since deforestation is 
already widespread on Salibabu and in particular Kabaruan, it would 
be interesting to assess the present distribution of enigma on these 
islands. A clearer indication of the species’ ability to adapt to secondary 
habitats, in competition with chloris, would then be available and 
predictions of the possible effect of logging on the species’ population 
would be easier to assess. 


Acknowledgements 


The authors would like to thank the following organisations who provided financial 
sponsorship for Action Sampiri: June Chamberlain Charity, British Petroleum, BirdLife 
International, Flora & Fauna International, Oriental Bird Club, The Parrot Society, 
Percy Sladen Memorial Fund, The University of York, Stiftung Avifauna Protecta, 
Royal Geographical Society and British Ornithologists’ Union. Research in Indonesia 
was conducted jointly with Universitas Sam Ratulangi, Manado, Sulawesi and under the 
sponsorship of Pusat Penelitian dan Pengembangan Biologi, Bogor and Lembaga Ilmu 
Pengatehuan Indonesia. 


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Meyer, A. B. & Wiglesworth, L. W. 1895. Eine zweite Sammlung von Vogeln von den 
Talaut-Inseln. Abh. Ber. K. Zool. Mus. Dresden (1894/95) 9: 1-9. 

Meyer, A. B. & Wiglesworth, L. W. 1898. The Birds of Celebes and Neighbouring Islands. 
Friedlander, Berlin. 

Oberholser, H. C. 1919. A revision of the subspecies of the White-collared Kingfisher, 
Sauropatis chloris (Boddaert). Proc. U.S. Nat. Mus. 55: 351-395. 

PPA 1980. Telaah Kemungkinan Pengembangan Taman Buru P. Karakelang, Sulawesi 
Utara. PPA, Bogor. 

Sibley, C. G. & Monroe, B. L. Jr. 1990. Distribution and Taxonomy of Birds of the World. 
Yale University Press, New Haven and London. 

Taylor, J. 1991. Recce report of ““Sangir-Talaud’? Oct-Nov 1990. Unpublished report. 

White, C. M. N. & Bruce, M. D. 1986. The Birds of Wallacea (Sulawesi, The Moluccas 
& Lesser Sunda Islands, Indonesia. B.O.U. Check-list No. 7, British Ornithologists’ 
Union, Tring. 


Addresses: Jon Riley, 1 The Croft, North Sunderland, Seahouses, Northumberland 
NE68 7XA, U.K. Damien Hicks, Little Wyllies, Deakes Lane, Cuckfield, West 
Sussex RH175JB, U.K. James C. Wardill, 27 Carlton Avenue, Hornsea, East 
Yorkshire HU18 1JG, U.K. 


© British Ornithologists’ Club 1998 


In Brief 118 . Bull. B.O.C. 1998 198Q) 
A hybrid munia? 


by S. (Bas) van Balen 
Received 21 April 1997 


In Bull. Brit. Orn. Cl. 116: 137-142, R. L. Restall described in detail 
the new munia Lonchura pallidiventer. The bird has a very distinctive 
appearance and seems to make a good (and attractive) new species. 
However, I feel that the author did not convincingly exclude the 
possibility of a hybrid in suggesting the species’ authenticity. The 
following incongruities were found in his account. 

1. The birds described show a suspicious amount of variation in 
plumage pattern, and the following characters in particular were 
apparently not consistent and not linked with sex: (a) presence or 
absence of black tips to the under tail coverts; (b) presence or absence 
of white spots on the breast; (c) colouring of heel barring: 
chestnut/black or buff/black; (d) colour of lower breast: black or 
chestnut. 

2. It appears that no offspring were produced by the birds, some of 
which were reportedly kept in captivity by apparently expert munia 
breeders for five years. ‘Though munias are not always prolific breeders, 
and some even reputedly difficult, this low fertility (or perhaps sterility) 
of the birds, suggests hybrids. 

3. ‘Though hardly any part of Kalimantan has been explored very 
exhaustively, the hinterland of Banjarmasin is one of the best surveyed 
areas, and new species are quite unexpected from this region. 

Whilst looking at the illustrations of Scaly-breasted Munia Lonchura 
punctulata and White-bellied Munia Lonchura leucogastra (both 
depicted very conveniently on one plate by Clement et al. 1993) I could 
not help thinking of Cream-bellied Munia being the perfect cross of 
these two species. The southern Kalimantan race of White-bellied, 
castanota, is very distinct from the other races because of its chestnut or 
deep warm brown upper parts, contributing even more to the rich 
brown uppers of Cream-bellied. Moreover, both species co-occur in 
South Kalimantan and are scarce (Smythies 1981; Holmes & Burton 
1987), which would promote hybridization as choice of mates is 
restricted (see Campbell & Lack 1985). The Cream-bellied’s slightly 
larger size than either of these species can be explained by hybrid 
vigour producing larger and stronger birds (Campbell & Lack 1985). 

Two sex-linked differences were described by Restall: (1) the 
grizzled lines on the upper part of the cream belly, which are also 
found in both sexes of White-bellied; (2) slight barring on the lower 
rump, which is found in both sexes of Scaly-breasted. ‘This may be 
consistent with subtle sexual differences found in both, supposedly 
monochromatic ‘parent’ species. 

At least three things should be done to ‘test’ this new species. 
1. Breed the new species and examine its fertility and consistency of 
specific characters in their offspring, 2. Cross-breed White-bellied and 
Scaly-breasted Munias, 3. Find the birds in the wild. Especially the last 


A. Townsend Peterson 119 Bull. B.O.C. 1998 118(2) 


is of importance because it remains questionable whether or not these 
birds, if hybrids, are of a natural provenance or a product of genetic 
manipulation by a local aviculturist. High prices are sometimes paid for 
rare, exotic bird species, and it is conceivable that especially new and 
attractive-looking forms would make good prices on the national and 
international market. The relatively large number of birds during the 
last five years (13 specimens), as reported by the author, suggests the 
latter possibility. 


References: 

Campbell, B. & Lack, E. (eds) 1985. Dictionary of Birds. 'T. & A. D. Poyser, Calton. 

Clement, P., Harris, A. & Davis, J. 1993. Finches and Sparrows. Helm, London. 

Holmes, D. A. & Burton, K. D. 1987. Recent notes on the avifauna of Kalimantan. 
Kukila 3: 2-32. 

Smythies, B. E. 1981. The Birds of Borneo. The Sabah Society with The Malayan Nature 
Society. 


Address: Birdlife International—Indonesia Programme, P.O. Box 310/Boo, Bogor 16003, 
Indonesia. 


© British Ornithologists’ Club 1998 


The distribution and type locality of the 
extinct Slender-billed Grackle Quiscalus 
palustris 


by A. Townsend Peterson 
Received 3 Fune 1997 


The Slender-billed Grackle Quiscalus palustris was described by 
Swainson (1827), based on specimens collected by W. Bullock. The 
type locality was described as ‘“‘marshes and borders of the lakes round 
Mexico”’, which was long interpreted as indicating the large lakes that 
formerly existed within the Valley of Mexico itself. However, this 
interpretation, and indeed the type locality of the species itself, were 
changed by Dickerman (1965), and accepted by most subsequent 
authorities (e.g. American Ornithologists’ Union 1983). 
Dickerman (1965) provided convincing evidence that the species also 
occurred in the marshes east of the Valley of Mexico along the 
headwaters of the Rio Lerma. E. A. Goldman collected the species at 
Lerma in 1904, describing the habitat as follows: ‘The marsh is filled 
with a varied assortment of aquatic vegetation, including tules, sedges, 
and many submerged species ... The marsh is an important watering 
place for migratory w aterfowl, and a breeding area for resident 
waterfowl” (Goldman 1951). Wilmot W. Brown, Jr., collected 
additional series at ‘““San Mateo” in 1910. Dickerman (1965) argued 
that this locality also was in the Lerma marshes; his identification of 
this site as San Mateo Atenco, 13.5 km ESE of Toluca, is borne out by 


A. Townsend Peterson 120 Bull. B.O.C. 1998 118(2) 


localities given on a few of Brown’s labels, which read ‘‘Mexico, Toluca 
(13.5 km ESE; San Mateo)’. Hence, the occurrence of Slender-billed 
Grackles in the Lerma marshes is well established (Dickerman 1965, 
Hardy 1967). 

However, Dickerman (1965) went on to argue that the species never 
occurred in the Valley of Mexico, and that in fact no grackle had 
occurred there until the 1950s, when a population of Great-tailed 
Grackles Q. mexicanus was established in Xochimilco. Nevertheless, 
the marsh habitats in the Valley of Mexico were of impressive 
dimensions. In 1520, Hernando Cortés described his first view of the 
valley, in the vicinity of Ixtapalapa, as follows (Morris 1928): 


‘There are trees and flowering shrubs, and with the lake there are innumerable fish and 
birds, such as wild duck, wigeon, and other waterfowl, and in such number that they 
almost cover the surface of the water. 


Descriptions provided by several other sixteenth century visitors to the 
Valley of Mexico (e.g. Gage 1980) support Cortés’ view of its immense 
marshes. After 300 years of European presence, however, in the early 
1800s, the valley was described by Mme. Calderon de la Barca (1987; 


translation mine) as follows: 


‘The scenery on this side of Mexico is arid and flat, and where the waters of the 
Lagunas, covered with their gay canoes, once surrounded the city, forming canals 
through its streets, we now see melancholy marshy lands, little enlivened by great 
flights of wild duck and waterfowl. 


Clearly, these marshes would originally have constituted ideal habitat 
for Slender-billed. Grackles, much as in the case of the Nicaraguan 
Grackle Q. nicaraguensis and Lake Nicaragua, but were on their way to 
extinction by the mid-nineteenth century. 

The type locality provided by Swainson (1827) indicates that the 
type material was indeed taken in the Valley of Mexico. In the same 
contribution, Swainson reported on 65 species taken in Mexico, at sites 
including ‘“‘near Vera Cruz’’, “Table land’’, Real del Monte, 
Temiscaltipec [sc], ““Sides of the Cordilleras’’, and “‘maritime land”’; 
one other specimen was described as having been purchased in the city 
of Mexico. Only that of the Slender-billed Grackle was referred to as 
‘“‘the lakes round Mexico’’, suggesting that the material was not just 
another haphazardly labelled record from ‘Temascaltepec or en route to 
or from. Remembering that in Mexico, the term “‘Mexico’”’ frequently 
refers to the city, Swainson’s type locality could easily refer to the lakes 
and marshes of the Valley of Mexico. 

Apart from the type specimen, however, other records exist of 
Slender-billed Grackles from the Valley of Mexico. Herrera (1891; 
translation mine), in a report on the vertebrates of the Valley of 
Mexico, listed the occurrence of Slender-billed Grackles as follows: 


In the marshy areas, among mammals, Arvicola pinetorum, A. mexicana, and Mustela 
brasiliensis; among birds, besides cranes and waterfowl, Pandion haliaetus (Fishing 
Hawk), Ceryle alcyon, Ceryle cabanisi (kingfisher), Sayornis nigricans, Sturnella magna 
mexicana, Anthus ludovicianus, Quiscalus tenuirostris [=Q. palustris], Melospiza fasciata 
mexicana, Cinclus mexicanus, Anthus ludovicianus, Cistothorus palustris ... 


A. Townsend Peterson 121 Bull. B.O.C. 1998 118(2) 


Also, Herrera (1889; translation mine) made the following footnote to a 
list of the vertebrates of the Valley of Mexico: 


New research has confirmed what I have already said about this species being confined 
to Xochimilco. According to the Biologia [Salvin and Godman’s Biologia 
Centrali-Americana], it is not Quiscalus macrourus [=Q. mexicanus], but QO. tenuirostris 
Sw., or Scaphidurus palustris Sw. 

The grackles of Xochimilco prefer to feed on corn, which is as abundant there as 
elsewhere in the valley. 


Hence, given reports of the species in the scientific literature and the 
abundant available habitat, the former occurrence of Slender-billed 
Grackles in Xochimilco, at the southern end of the Valley of Mexico, 
seems certain, and the type locality is probably correctly interpreted as 
referring to the Valley of Mexico. 

Since the arrival of Europeans, however, the great lakes of the Valley 
of Mexico have been severely degraded. Worst of all, between 1609 and 
the first part of the present century, a series of canals was cut to drain 
the lakes, and the marsh habitats there have been destroyed completely. 
This destruction probably led to the species’ extinction in the Valley of 
Mexico prior to the twentieth century, leaving only the Lerma 
populations extant when Goldman and Brown were collecting. The 
Lerma populations probably did not last much longer than 1910, as no 
substantially later records are known. 


Acknowledgements 


I thank Richard C. Banks for sending a copy of the original description of the species, and 
Rosa M. Salazar for amiable company during hours of research in the library. 


References: 

American Ornithologists’ Union. 1983. Check-list of North American Birds. 6th edn. 
American Ornithologists’ Union. 

Calderon de la Barca. 1931. Life in Mexico, During a Residence of Two Years in that 
Country. Mexico Press, Mexico 

Dickerman, R. W. 1965. The juvenal plumage and distribution of Cassidix palustris 
(Swainson). Auk 82: 268-270. 

Gage, T. 1980. Viajes en la Nueva Espana. Casa de las Américas, La Habana. 

Goldman, E. A. 1951. Biological investigations in Mexico. Smithsonian Misc. Coll. 115: 
1-476. 

Hardy, J. W. 1967. Evolutionary and ecological relationships between three species of 
blackbirds (Icteridae) in central Mexico. Evolution 21: 196-197. 

Herrera, A. L. Jr. 1889. Notas acerca de los vertebrados del Valle de México. La 
Naturaleza, 2nd ser., 1: 299-342. 

Herrera, A. L. Jr. 1891. El clima del Valle de México y la biologia de los vertebrados. La 
Naturaleza, 2nd ser., 2: 38-86. 

Morris, J. B. (ed.) 1928. Hernando Cortés: Five Letters, 1519-1526. W. W. Norton Co., 
Inc., New York. 

Swainson, W. 1827. A synopsis of the birds discovered in Mexico by W. Bullock, F.L.S. 
and H.S., and Mr. William Bullock, jun. Philosoph. Mag., new ser., 1: 433-442. 


Address: Natural History Museum, The University of Kansas, Lawrence, Kansas 66045, 
U.S.A. 


© British Ornithologists’ Club 1998 


In Brief 122 Bull. B.O.C. 1998 118(2) 


First record of the Striped Manakin 
Machaeropterus regulus in Guyana 


by David Ff. Agro © Robert S. Ridgely 


Received 6 Fune 1997 


While examining specimens in the collection of the Academy of 
Natural Sciences of Philadelphia (ANSP), we were surprised to find a 
specimen of Machaeropterus regulus that, for over 100 years, had been 
incorrectly identified as a Tiny ‘Tyrant-Manakin Tyranneutes virescens. 
This specimen represents the first and, to our knowledge, only record 
of M. regulus for Guyana. 

The Striped Manakin Machaeropterus regulus is an uncommon to 
locally fairly common bird of humid forest and mature secondary 
woodland in much of northern and northwestern South America; an 
isolated population occurs in eastern Brazil (Ridgely & Tudor 1994). 
The species has not been recorded in northeastern South America (e.g. 
no records from Guyana: Chubb 1921, Snyder 1966; Surinam: 
Haverschmidt & Mees 1994; French Guiana: Tostain et al. 1992). 
Apart from the isolated Brazilian population, the previous easternmost 
record was from Rio Suruktn (La Faisca) at the headwaters of the Rio 
Caroni in southern Bolivar, Venezuela (Phelps & Phelps 1950). 

The specimen (ANSP 51442) is a female collected by Henry Whitely 
Jr. (1844-1893) on 10 October 1890 near Ourumee in what was then 
British Guiana. Ourumee is another name for the Merumé Mountains, 
5°48'N; 60°6'W (Stephens & Taylor 1985). Based on range, we have 
tentatively assigned this specimen to M. r. aureopectus, a subspecies 
described by Phelps Gillard (1941) on the basis of a single male from 
Kabadiscana in southern Bolivar, Venezuela. 

Although M. regulus is difficult to see, it should be easily detected in 
the field by its calls. A number of observers (RSR, Mark, B. Robbins, 
Tristan J. Davis, Davis Finch), who are familiar with these calls, have 
visited potentially suitable areas for M. regulus in Guyana over the past 
five years without reporting this species. This suggests that M. regulus 
is local and rare in Guyana, but observers should be aware that it could 
occur elsewhere in the Guianas. 


This observation was made in connection with our recent fieldwork in Guyana for the 
Iwokrama Faunal Survey made possible through the support of the Government of 
Guyana, the Iwokrama International Centre for Rain Forest Conservation and 
Development, the United Nations Development Programme, and the University of 
Guyana. We thank Graham G. Watkins for his help in coordinating the Academy’s field 
activities in Guyana, Vicki Funk and the Smithsonian Institution’s Biodiversity of the 
Guianas Program for its support in Georgetown, and Sally B. Conyne and Leo Joseph for 
their suggestions in preparing the manuscript. 


References: 

Chubb, C. 1921. The Birds of British Guiana. British Museum, London. 

Phelps, W. H. & Gilliard, E. T. 1941. Seventeen new birds from Venezuela. dm. Mus. 
Novit. no. 1153. ; 

Phelps, W. H. & Phelps, W. H., Jr. 1950. Lista de las aves de Venezuela con su 


distribucién; Parte 2, Passeriformes. Caracas. 


In Brief 123 Bull. B.O.C. 1998 118(2) 
Haverschmidt, F. & Mees, G. F. 1994. Birds of Surinam. VACO N.V. Uitgerversmaat- 


schappij, Paramaribo, Surinam. 

Ridgely, R. S. & Tudor, G. 1994. The Birds of South America. Vol. 2. Oxford University 
Press. 

Snyder, D. E. 1966. The Birds of Guyana. Peabody Museum, Salem, Massachusetts. 

Stephens, L. & Traylor, M. A., Jr. 1985. Ornithological Gazetteer of the Guianas. 
Harvard University, Cambridge, Massachusetts. 

Tostain, O., Dujardin, J.-L., Erard, C. & Thiollay, J.-M. 1992. Oiseaux de Guyane. 
Muséum National d’Histoire Naturelle, Paris. 


Address: The Academy of Natural Sciences, Department of Ornithology, 1900 Benjamin 
Franklin Parkway, Philadelphia, PA 19103, U.S.A. 


© British Ornithologists’ Club 1998 


The first documented nest and eggs of Merops 
muellert mentalis 


by Michael F. Carter & Robert W. Dickerman 
Received 8 Fune 1997 


Gatter (1988) noted that the Blue-headed Bee-eater Merops muelleri 
mentalis is a resident breeder of Liberia. That status was assumed since 
the nests or eggs of the Upper Guinea subspecies had never been 
found. Gatter (1988) also listed M. m. mentalis in a category for which 
all possible information about the species in Liberia is needed. 

Nests and eggs of the Lower Guinea subspecies, M. m. mueller1, have 
been found on three occasions (Bannerman 1955, Fry 1984, Fry et al. 
1992). These nests were described as 55-60 cm deep burrows with 
terminal, oval, unlined egg chambers. Nests were found in a roadside 
bank, in the wall of a sawyer pit in Cameroon, and along a forest path in 
Gabon. One nest from Cameroon contained eggs in January, and 
nestlings in February (Fry et al. 1992). In Gabon, nest building began 
in October and young fledged in February. Eggs from Cameroon were 
described as white and subspherical, measuring 23.2 < 20.1 and 
23.7 X 20.0 mm (Fry et al. 1988). 

M.m. mentalis was first recorded in Liberia at Mount Nimba by 
Colston & Curry-Lindahl (1965). During their studies 10 specimens 
were obtained, none were in breeding condition, but females in August 
and September with ovaries measuring 7mm might have been 
approaching breeding condition. Birds captured in July were in the 
middle to late stages of wing moult, indicating they might have just 
finished nesting. 

A nest of Merops muellert mentalis was found on 8 March 1990 by 
MFC while walking along a shallow, mostly dry stream bed. The 
stream bank was c. 30 cm tall and was probably within 3 km of shallow 
running water but none was seen in the immediate area. The habitat 
was primary semi-deciduous rainforest with a canopy approaching 
25 m in height and a dense growth of understory trees near the stream 


In Brief 124 Bull. B.O.C. 1998 118(2) 


bed. The nest locality was c.11km north of Zigida (Lofa County) at 
540 m in the Lofa Mountains of northern Liberia. 

‘The nest burrow was in a gently sloping portion of a sandy bank 
c. 45 cm above a small pool of water. It was c. 40 cm deep with a 45 mm 
opening. A bird flew from the burrow and perched in the lower limbs 
of an understory tree allowing close observation and positive 
identification. The two eggs were subspherical, cream-white (speckled 
brown from earth), and both measured 22.5 <x 18.0 mm; they were 
unincubated. They have been deposited in the American Museum of 
Natural History, New York. 

Details accord closely with nests and eggs of the nominate race, but 
the Liberian nest was in a gently sloping (not steep) bank, and its 
burrow was shallower. Nests of most bee-eater species vary in burrow 
depth and site characteristics (Fry et al. 1992). 


We thank Alexander Peal, head of Wildlife & National Parks of the Forestry 
Development Authority (Liberia), for assistance during the course of this work. Funding 
was provided by National Geographic Society and American Museum of Natural 
History. 


References: 

Bannerman, D. A. 1995. The Birds of West and Equatorial Africa. Vol. 1. Oliver and 
Boyd, Edinburgh. 

Colston, P. R. & Curry-Lindahl, K. 1986. The birds of Mount Nimba, Liberia. British 
Museum (Natural History), London. 

Fry, C. H. 1984. The Bee-eaters. Buteo Books, Vermillion, South Dakota. 

Fry, C. H., Keith, S. & Urban, E. K. (eds) 1988. The Birds of Africa, Vol. 3. Academic 
Press. 

Fry, C. H. & Fry, K. 1992. Kingfishers, Bee-eaters and Rollers, A Handbook. Princeton 
Univ. Press. 

Gatter, W. 1988. The birds of Liberia (West Africa), a preliminary list with status and 
open questions. Verh. Orn. Ges. Bayern 24: 689-723. 


Addresses: M.F.C.—Colorado Bird Observatory, 13401 Piccadilly Road, Brighton, 
Colorado 80601, U.S.A. R.W.D.—Biology Department, Museum of Southwestern 
Biology, Albuquerque, New Mexico 87131, U.S.A. 


© British Ornithologists’ Club 1998 


Monotypy of Francolinus griseostriatus 


by N. F. Collar 
Received 20 Fune 1997 


The Grey-striped Francolin Francolinus griseostriatus is one of a suite 
of species endemic to the escarpment zone of western Angola, where 
it occurs in two apparently disjunct populations, a northern one 
extending from Cuanza Norte and Malanje to Cuanza Sul, and a 
southern one confined (after a boundary extension that embraced 
northern Huila) to the province of Benguela (Hall 1961, 1963, Pinto 
1983, Urban et al. 1986). 


In Brief 125 Bull. B.O.C. 1998 118(2) 


Pinto (1983) treated these two populations as racially distinct, based 
on eight specimens from Benguela (in the Instituto de Investigacao 
Cientifica de Angola, Luanda) having generally darker underparts, with 
more cinnamon than buff (“‘fulvo-esbranquicado’’) shading to the 
feather edges and broader, deeper brown feather centres, and with 
undertail coverts marked with darker spots and bars. He also detected a 
very slight increase in size in these birds, albeit with much overlap. 
However, he felt unable to give them a name for want of immediate 
comparative material. 

Ironically, outside Angola it is the want of southern birds in 
collections that has delayed progress in this matter, but one was located 
in the Carnegie Museum of Natural History (CM), Pittsburgh, and 
another proves to exist in the series of six held by the American 
Museum of Natural History (AMNH), New York. In May 1997 the 
Carnegie bird, CM 108689, a female from Chingoroi, Benguela, 
13°37’'S 14°01’E, taken at 670 m in October 1930 by R. and L. Boulton, 
was sent to New York for examination alongside the five adult AMNH 
specimens (541414, female, Canhoca, 9°15’S 14°41’E, December 1903, 
by W. J. Ansorge; 348799, male, Ndala Tando, 9°18’S 14°54’E, 
undated, no collector; 541410, female, and 541411, male, Ndala Tando, 
September 1908, by W. J. Ansorge; 541413, male, Bongo River, 
Benguela, 13°25’S 14°40’E, August 1904, by W. J. Ansorge). All 
coordinates are from Traylor (1963). 

From above, no differences in these six birds are apparent that 
would not be attributable to minor individual variation. From below, 
the two most distinct are AMNH 541414 and 348799, which have 
the tawny markings on the breast, flanks and belly reduced to fairly 
narrow streaking. By contrast, 541410 and 541411 possess substan- 
tially broader, blotch-like markings on the breast. However, the last 
three are from the same locality, and the first is from a site not far 
distant, suggesting that either individual or age-related factors are in 
play. 

More significantly, 541410 and 541411 are little if at all different 
from 541413 and CM 108689. It is true that the tawny markings on 
the undersides of the CM bird appear very slightly broader and 
more extensive than on the other specimens, most notably 541410; 
but the differences between 541413 and 541411 are entirely 
negligible and militate heavily against any racial separation of the 
two populations they represent. Nothing in the shading of the 
markings on the underparts or on the undertail coverts on any of 
the specimens indicated any consistent differences between the 
representatives of the two populations. No measurements were taken, 
although wing-lengths were qualitatively compared and found to be 
very similar. 

If heavier marking on the undersides is a sign of greater age, then the 
only way in which Pinto’s view might yet prevail is if AMNH 541410 
and 541411 are old birds showing the broadest feather blotching that 
representatives of the northern population ever achieve, while 541413 
and CM 108689 are young birds yet to acquire the distinctiveness that 
Pinto recognised in the Luanda series. This seems rather improbable, 


In Brief 126 Bull. B.O.C. 1998 118(2) 


however, and certainly for the time being Francolinus griseostriatus 
must remain monotypic. 


Robin Panza at Carnegie Museum arranged the loan of the CM specimen to New York, 
and Paul Sweet and Christine Blake of the American Museum of Natural History 
respectively allowed access to material there and returned the loan; Pamela Rasmussen 
also examined the material and concurred with the judgement expressed above. Their 
kindness is deeply appreciated. 


References: 

Hall, B. P: 1961. The faunistic importance of the Scarp of Angola. [bis 102: 420-442. 

Hall, B.P. 1963. The francolins, a study in speciation. Bull. Brit. Mus. Nat. Hist. (Zool.) 
10: 105-204. 

Pinto, A. A. da R. 1983. Ornitologia de Angola, 1. Instituto de Investigacao Cientifica 
Tropical, Lisboa. 

Traylor, M. A. 1963. Check-list of Angolan Birds. Publ. Cult. Co. Diam., Angola, no. 61. 

Urban E. K., Fry, C. H. & Keith, S. (eds) 1986. The Birds of Africa. Vol. 2. Academic 
Press, London. 


Address: BirdLife International, Wellbrook Court, Girton Road, Cambridge CB3 0NA, 
WK: 


© British Ornithologists’ Club 1998 


The correct citation of Coragyps (Cathartinae) 
and Ardeotis (Otididae) 


by Steven M.S. Gregory 
Received 2 October 1997 


Coragyps (Cathartinae) and Ardeotis (Otididae) were first used and 
established in the same work, Le Maout (1853). Coragyps appears on 
pp. 57 (in key) and 66, Ardeotis on pp. 339 (in key) and 340. However, 
Coragyps is ascribed to Geoffroy Saint-Hilaire, by Gray (1855), Peters 
(1931) and Stresemann & Amadon (in Peters 1979), while Ardeotis is 
ascribed to Le Maout, by Meinertzhagen (1954) and Del Hoyo et al. 
(1996). Clearly, one of these citations is in error. Coragyps is a well 
established genus and it is unlikely that any recent author would 
question the usual citation. Stresemann & Amadon (in Peters 1979) 
appear not to have looked at the original reference, but relied on 
seemingly well-founded previous designations. Thus they refer to 
‘Geoffroy Saint-Hilaire’ as being ‘in Le Maout’, which implies that 
Geoffroy Saint-Hilaire wrote that particular section of the book, 
whereas Le Maout was quoting from unpublished material. Ardeotis, 
on the other hand, is entirely absent from all of the standard references, 
and its replacement of Choriotis G. R. Gray (1855) is poorly 
documented. 

It is therefore proposed that the citation for Coragyps should be in 
agreement with that for Ardeotis and that they should read: 


Coragyps Le Maout, 1853, Histoire Naturelle des Oiseaux suivant la 
Classification de M. Isidore Geoffroy Saint-Hilaire, pp. 57 (in key), 66. 


In Brief 127 Bull. B.O.C. 1998 118(2) 


Type by monotypy, Coragyps urubu “‘Isid. Geoffroy’ = Vultur atratus 
‘Wilson’ Bechstein. 


Ardeotis Le Maout, 1853, Histoire Naturelle des Oiseaux suivant la 
Classification de M. Isidore Geoffroy Saint-Hilaire, pp. 339 (in key), 
340. Type by monotypy, Otis arabs Linnaeus. 


Acknowledgements 


I should like to thank Michael Walters, NHM, Tring, Dr. Alan Knox and Dr. David 
Snow for their generous help. 


References: 

Del Hoyo, J., Elliot, A. & Sargatal, J. (eds) 1996. Handbook of the Birds of the World, 
vol. 3. Lynx Edicions, Barcelona. 

Gray, G. R. 1855. Catalogue of the Genera and Subgenera of Birds. British Museum, 
London. 

*Le Maout, E. (1853) 1855. Histoire Naturelle des Oiseaux suivant la Classification de M. 
Isidore Geoffroy Saint-Hilaire. L. Curmer, Paris. 

Meinertzhagen, R. 1954. Birds of Arabia. Oliver & Boyd, Edinburgh. 

Peters, J. L. 1931. Check-list of Birds of the World, vol. 1. Harvard University Press, 
Cambridge, Mass. 

Stresemann, E. & Amadon, D. 1979. In Check-list of Birds of the World, vol. 1., 2nd 
edition. Museum of Comparative Zoology, Cambridge, Mass. 


*=Examined by Michael Walters. 
Address: 35 Monarch Road, Northampton, Northamptonshire NN2 6EH, U.K. 
© British Ornithologists’ Club 1998 


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NOTICE TO CONTRIBUTORS 

Papers are invited from Club Members or non-members, especially on 
taxonomic and distributional topics; descriptions of new species are especially 
welcome and may be accompanied by colour photographs. Two copies of 
manuscripts, typed on one side of the paper, double spaced and with wide 
margins, should be sent to the Editor, Prof. Chris Feare, 2 North View 
Cottages, Grayswood Common, Haslemere, Surrey GU27 2DN, UK. All 
contributions, including In Brief articles, should follow the style of main 
papers in this issue of the Bulletin. 


A contributor is entitled to 10 free offprints (16 if 2 or more authors) of the pages of the 
Bulletin in which his contribution, if one page or more in length, appears. Additional 
offprints or offprints of contributions of less than one page may be ordered when the 
manuscript is submitted and will be charged for. Authors may be charged for proof 
corrections for which they are responsible. 


MEMBERSHIP 

Only Members of the British Ornithologists’ Union are eligible to join the 
Club, and to receive (postage free) four quarterly issues of the Bulletin, and the 
annual index, for an annual subscription of £12 (or U.S. $26). Applications, 
enclosing the annual subscription, should be made to the Hon. Secretary 
(address as below). 

The 1998 List of Members, and addresses will not be published with the 
Bulletin this year, but copies are available, on applicaticn (with a remittance 
of £1.00 to cover costs of production and postage), to the Hon. Secretary. 
Please advise the Hon. Secretary, without delay, of any address changes, or 
corrections, for despatch of the Bulletin. 


NON-MEMBER SUBSCRIBERS & APPLICATIONS FOR BACK 
NUMBERS OR OTHER PUBLICATIONS 

The Bulletin (for 1998 onwards), together with annual index, may be 
purchased (postage free) by Non-member Subscibers on payment of an annual 
subscription of £22 (or U.S. $45) on application to The Publications Officer, 
S. J. Farnsworth, Hammerkop, Frogmill, Hurley, Maidenhead, Berks SL6 5NL, 
U.K. Single issues, and runs of back numbers of the Bulletin, and also other 
BOC Publications may similarly be obtained, on request to him. 


PAYMENTS 

All amounts quoted are net and should be paid in £ sterling, if possible. 
Payments in other currencies must include a further £4 for UK bank charges 
(except for annual rates in U.S. dollars, which are inclusive). All cheques or 
drafts should be made payable to the British Ornithologists’ Club. If preferred, 
remittances may be made by bank transfer direct to the Club’s bank 
account—Barclays Prime Account, Dale House, Wavertree Boulevard, 
Liverpool L7 9PQ, U.K. (Sort Code 20-00-87 Account No. 10211540), with 
confirmation to the Hon. Treasurer, D. J. Montier, Eyebrook, Oldfield Road, 
Bickley, Bromley, Kent BR1 2LF. 


CORRESPONDENCE 
Correspondence on membership, changes of address and all other matters 
should be addressed to the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA, U.K. For details 
of Club Meetings see inside front cover. 


Registered Charity No. 279583 


CONTENTS 


LAMBERT, F. R. A new species of Amaurornis from the Talaud Islands, 
Indonesia, and a review of taxonomy of bush hens occurring from the 
Philippines to Australasia 2.052... )00. . 24 os Yas oe 

ZYSKOWSKI, K., PETERSON, A. T. & KLUZA, D. A. Courtship behaviour, vocaliz- 
ations, and species limits in Atthis hummingbirds ................ 

GAMAUF, A. & PRELEUTHNER, M. A new taxon of the Barred Honeybuzzard 
Pernis celebensis from the Philippines. ...:........5.02 eee 

DELESTRADE, A. Distribution and status of the Ethiopian population of the 
Chough Pyrrhocorax pyrrhocorax baileyi .............0)eeneeee 

DEMEY, R. & FISHPOOL, L. D.c. On the existence of a melanistic morph of the 
Long-tailed Hawk Urotriorchis macrourus ...........2..+e+e+00e08 

COLLINS, C.T. Food delivery and chick provisioning in the cypscloidine swifts 

RILEY, J., HICKS, D. & WARDILL, J.c. The taxonomic status of Halcyon enigma 
on the Talaud Islands, Indonesia ................. Joe 

In Brief van BALEN, S. A hybrid munia? ...............5 =e 

PETERSON, A. T. ‘The distribution and type locality of the extinct 
Slender-billed Grackle Quiscalus palustris ............... 
AGRO, D. J. & RIDGELY, R. S. First record of the Striped Manakin 
Machaeropterus regulus in Guyana...........+. a2s2 eee 
CARTER, M. F. & DICKERMAN, R. W. ‘The first documented nest and 
eggs of Merops muellert mentalis.... 5. ........ 0 
COLLAR, N. J. Monotypy of Francolinus griseostriatus.......... 
GREGORY, S. M.S. The correct citation of Coragyps (Cathartinae) 
and Ardeotis (Otidae) ...... 02.6. 05. be. | ee) ee 


101 


105 
108 


113 
118 


119 


122 


123 
124 


126 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 


mail, after their current subscription has been paid. 


COMMITTEE 


Revd T. W. Gladwin (Chairman) (1997) Dr R. P. Prys-Jones (1995) 
Mrs A. M. Moore (Vice-Chairman) (1997) N. H. F. Stone (1995) 
Cdr M. B. Casement, OBE, RN D. Griffin (1997) 
(Hon. Secretary) (1996) N. J. Redman (1997) 
D. J. Montier (Hon. Treasurer) (1997) Mr R. E. Scott (1998) 


Hon. Editor: Prof C. J. Feare 
Chairman of Publications Sub-committee: Dr R. P. Prys-Jones 
Publications Officer: S. J. Farnsworth 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


AL 
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eec- ISSN 0007-1595 
Bi. RD 


Bulletin of the 


British Ornithologists’ Club 


Edited by 
Prof CHRIS FEARE 


\ 


OCT 2 9 996 


LIBRARIES 


Pe ae 


Volume 118 No. 377"” September 1998 


FORTHCOMING MEETINGS 


Advance notice of meeting dates for 1999. Eight meetings have been arranged for the 
following Tuesdays: 19 January (K. F. Betton on “Birding by Ear’’—see below), 16 March, 
20 April, 4 May (AGM and social evening—with informal “mini-talks’”), 6 July, 
7 September, 12 October and 30 November. Details of speakers on these dates will be 
published when finalised. 


Tuesday 17 November 1998. Dr W. G. (Bill) Porteous will speak on “Birds of the 
Humboldt Current’. Bill was born and raised in Shetland, and may therefore be unique in 
having added Booted Warbler to his life list before he added Blue Tit. He subsequently 
qualified as a geologist, and this has given him the opportunity to pursue birds in various parts 
of the world, particularly in north and south America. He recently spent five years in Colombia, 
which provided some welcome exposure to the neotropics, which had long been a particular 
interest of his. The opportunity to participate in a voyage southwards along the coasts of Peru 
and Chile in late 1995 provided the material we are to see this evening. 

Applications to the Hon. Secretary by 3 November, please. 


Tuesday 19 January 1999. Keith Betton will speak on “Birding by ear—a look at the 
world of strange, and not so strange bird sounds’. Keith was born and educated in 
London, where he quickly established a life-long interest in ornithology. He came to 
prominence when, at the early age of 12, he was appointed a member of the Department of 
Environment Committee on Bird Sanctuaries in Royal Parks, and was Official Observer for 
Bushy Park (1973-79). He has served on numerous ornithological committees, including 
Chairman Ornithology Section of London NHS (1984-92), President London NHS (1982-84), 
BTO Regional Representative Greater London (1981-93), and Council BTO (1987-91), also 
Committee BOC (1985-88). He is a prolific contributor to and editor of several journals. His 
high profile post as Head of Corporate Affairs, Association of British Travel Agents, involves 
travelling worldwide on business, but this now leaves him less time for these ornithological 
activities. His special interest is in Africa, and in sound recordings, of which he has 
accumulated a large library of his own. 

Applications to the Hon. Secretary by 5 January, please. 


Meetings are held in the Sherfield Building of Imperial College, South Kensington, London 
SW7. The nearest Tube station is at South Kensington, and car parking facilities are available; a 
map of the area will be sent to members, on request. The cash bar is open from 6.15 p.m., and 
a buffet supper, of two courses followed by coffee, is served at 7.00 p.m. (A vegetarian menu 
can be arranged if ordered at the time of booking.) Informal talks are given on completion, at 
about 8.00 p.m. 


Overseas Members visiting Britain are particularly welcome at meetings. For 
details in advance, please contact the Hon. Secretary, Cdr M. B. Casement, OBE, RN, 
Dene Cottage, West Harting, Petersfield, Hants GU31 5PA. 

Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1998 


Apart from single copies made for the purposes of research or private study, or criticism or 
review, as permitted under UK law, no part of this publication may be reproduced, stored 
or transmitted in any form or by any means, except with prior permission in writing of the 
publishers, or in accordance with the terms of licences issued by the Copyright Licensing 
Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; for 
address see inside back cover. 


Eee EEEOOeEeEeEeEEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEyeeee 


129 Bull. B.O.C. 1998 118(3) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 118 No. 3 Published September 23 


ANNUAL GENERAL MEETING 


The Annual General Meeting of the British Ornithologists’ Club was 
held in the Ante-room of the Sherfield Building, Imperial College, 
London SW7 on Tuesday 19 May 1998 at 6 p.m. with The Reverend 
T. W. Gladwin in the Chair. Apologies had been received from 
R. E. F. Peal and S. J. Farnsworth. 23 Members were present. 

The Minutes of the Annual General Meeting held on 20 May 1997, 
which had been published (Bull. Brit: Orn. Cl. 117: 153-156), were 
approved and signed by the Chairman. 


Chairman’s report. ‘The Chairman opened by saying that, although 
not customary for the Chairman to make a separate report, there were 
a number of interesting areas of Club activity this year, worthy of 
mention on this occasion. 

Last year, the Club had kindly elected him to succeed David Griffin 
as Chairman. Whilst recording his pleasure that the Committee could 
count on David's continued support and advice, as a member of the 
Committee, his name had been omitted in the minutes, as elected to 
replace Ronald Peal as Committee Member, in May 1997. He 
apologised to David for that oversight, which will be appropriately 
corrected in the record for this meeting. 

One of David Griffin’s many contributions, for which the Club is 
grateful, was to establish a journal in which he has set out details of all 
867 meetings, from 1892, through the period of his Chairmanship. The 
numbered record of each meeting includes details of venue, attendance, 
subject(s) and speaker(s). He personally, and he hoped future 
Chairmen, will maintain that journal, which will pass into the archives, 
when a second volume becomes necessary. 

The Club’s archives, kindly stored in the Natural History Museum 
at Tring, continue to be maintained and catalogued by Francis Stone. 
The Committee was particularly pleased when he agreed that his role 
should be acknowledged by appointing him as the Club’s first 
Archivist. It may be appropriate to identify and define this office in the 
Club’s Rules, at the next revision. The Committee had agreed that it is 
important that the Club’s transactions are properly recorded in the 
Bulletin, and have suggested that Francis should prepare an appropriate 
account of the archives. 

It has been customary for the Chairman to be custodian of a 
complete bound set of the Bulletins. The Committee has agreed that 
this set, currently held by himself, should be placed in the archives. 

By virtue of its encouragement and interest in systematics and 
taxonomy, the Club occupies an important niche in ornithology. In 


Meetings 130 Bull. B.O.C. 1998 118(3) 


making our contribution we have an inherent responsibility for all 
seriously proposed systems of biological classification to receive 
consideration. Since the publication of the centenary Volume 112A 
Avian Systematics and Taxonomy (Monk, J. F. 1992), several 
commentators have expressed regret at the relatively little attention 
given to the cladistic method of systematic classification. The cladistic 
approach, proposed by Hennig (Phylogenetic Systematics, 1966), has a 
significant number of supporters and it should be interesting, as well as 
desirable, to invite a speaker or paper on this subject. 

Whilst the Bulletin continues to attract a wealth of major papers, it is 
important that sufficient resources are made available for its continuing 
development, probably to include more colour plates, under the 
editorship of Prof. Chris Feare. 

The Club is fortunate in being endowed and trusted to fund 
scientifically and historically tmportant ornithological publications 
without, in every case, a primary concern for commercial viability. He 
believed that history will increasingly value the Club’s publications. 
The value of museum material in diagnosing the cause of egg-shell 
thinning in Peregrines Falco peregrinus, for example, illustrates the 
potential value to conservation of Avian Eggshells: an Atlas of Scanning 
Electron Micrographs (Mikhailov, K. E. 1997), published as BOC 
Occasional Publications No.2. The Club is grateful to Amberley 
Moore, who is handing the Chairmanship of the Publications 
Sub-Committee to Dr Robert Prys-Jones, to Prof. Chris Feare as 
Honorary Editor of the Bulletin, to John Farnsworth, Publications 
Officer, and to all other members of that Sub-Committee for the 
excellence and sales of Club publications. 

The Club enjoys warm relations with the Sub-Dept. of Ornithology 
of the National History Museum at Tring, which provides the facilities 
for storing our archives and stock of Bulletins. We are grateful to 
Robert Prys-Jones and his staff for this and other kindnesses, and 
especially to Mrs ‘Effe’ Warr, who has maintained the inventory of, 
and processes orders for, back numbers of the Bulletin. 'The Club 
continues to support the Museum, in whatever way may, from time to 
time, be possible. 

The Committee has started to think about how the Club might mark 
the Millennium, and would be pleased to receive suggestions from 
Members. i 

Several Members have questioned the authenticity of the Club’s 
gavel, which is said to have been made from wood from H.M.S. Beagle. 
Out of interest, he had started enquiries of a number of naval 
historians, to obtain details of H.M.S. Beagle’s disposal. 

Helen Baker, who now retires from the Committee, will continue to 
assist the Hon. Secretary with the membership records. The Club is 
grateful for her significant contribution. 

Finally, the Chairman recorded his thanks to all others who have 
contributed in so many ways to the administration and life of the Club, 
over the past year; to those already mentioned, to Michael Casement 
(Hon. Secretary), David Montier (Hon. Treasurer), all other Members 
of the Committee, the Trustees of the Herbert Stevens Trust Fund, to 


Meetings 131 Bull. B.O.C. 1998 118(3) 


Imperial College for the provision of parking and meeting facilities and 
valued dinners, and to the Members who have supported the excellent 
programme of meetings. 


The Annual Report of the Committee for 1997 had been 
published in Bull. Brit. Orn. Cl. 118(1): 1-3. 


The Bulletin. Prof. Chris Feare (Hon. Editor) reported that he had 
received an encouraging flow of papers for publication, at an average of 
four per month and, after the June issue, 35 are undergoing the process 
of preparation for publication. Issue 118(2) should appear on time, and 
would contain the description and a colour plate of a new species of rail, 
and he hoped the September issue would also contain a description of a 
new species, illustrated by a colour plate. A paper on a nomenclatural 
issue, published in the March issue 118(1) had stimulated some 
controversy. This highlighted the complexity of the rules governing 
nomenclature, and the diminishing number of people professionally 
employed as taxonomists, and who understand these rules. He would 
therefore be publishing the comments received from Storrs Olson. 

The Chairman thanked the Editor, on behalf of the Club, for his 
report, and his achievements, since taking over this post. This was 
enthusiastically endorsed by all present. 


The Annual Accounts for 1997 were presented by David Montier 
(Hon. Treasurer), who drew attention to the following points: 

Total assets at 31 December 1997 amounted to £258,173, which was 
some £32,000 higher than in the previous year. Most of this was 
attributable to a substantial increase in the value of the Herbert Stevens 
Trust fund investments, which rose by £29,300 over the year, 
reflecting a strong performance in stock markets, particularly in the 
U.K. 

The statement of Financial Activities shows a surplus of £2,383 on 
Unrestricted Funds over the year. Income on Unrestricted Funds rose 
by £4,900, the main components being about £1,600 increase in 
Members’ subscription income, £2,200 in investment income 
and £600 in sales of publications. The increase in subscription 
income was the result of the rise in the subscription rate, as from 
1 January 1997. 

Expenditure rose by almost £8,500. The major part of this was 
caused by the publication of two books in the same year: Burd 
Manuscripts and Drawings—(Occasional Publications No.2), at the 
beginning of the year, and Avian Eggshells—(Occasional Publications 
No. 3) at the end. The sum of £500 was also paid towards the 
production costs of the Uganda Atlas out of the Club’s general funds. 
Meeting costs had increased as a result of the introduction of a room 
hire charge by Imperial College. 

Adoption of the Accounts was proposed by Mrs Mary Muller, 
seconded by Dr David Snow, and approved by all present. 

The Chairman expressed the thanks of all to the Hon. Treasurer and 
the Trustees of the Herbert Stevens Trust Fund. 


Meetings 132 Bull. B.O.C. 1998 118(3) 


Publications Sub-Committee. Mrs Amberley Moore said that, as 
the Hon. Treasurer had just reported, two Occasional Publications, 
Nos 2 and 3, had been published, and work was in hand with two 
further projects: 

(i) A Bird Atlas of Uganda. 'This was now being progressed as a joint 
project with the Union, the final arrangements for which are under 
active discussion. The Club was grateful for a grant from the RSPB 
towards the work of preparing the distributional maps which is being 
done at the University of Makere in Uganda, in association with the 
East Africa Natural History Society. 

(11) Type Specimens of Birds in the University Museum of Zoology at 
Cambridge. At the beginning of the year the Club was approached by 
the University Museum of Zoology at Cambridge with a request to 
publish the catalogue of bird skins prepared by the late C. W. Benson, 
Editor of the Bulletin 1969-74. Negotiations with the Museum are 
proceeding, and it is hoped the catalogue will appear in print later this 
year. 

Further publication plans include the possibility of a monograph on 
some Mascarene birds by Storrs Olson et al., and the Proceedings of 
the joint BOU/BOC/NHM/BirdLife conference on museums and avian 
archives to be held in November 1999. The meeting expressed great 
satisfaction at progress with recent and proposed publications, and 
thanked Mrs Amberley Moore and her Sub-Committee accordingly. 


Election of Officers and Committee. The Chairman said that the 
Committee’s proposals had been published in Bulletin 118(1): 3, and no 
other nominations had been received. He proposed that Michael 
Casement be re-elected as Hon. Secretary, and David Montier be 
re-elected as Hon. Treasurer. This proposal was seconded by Mrs 
Mary Muller and approved by all present. 

He proposed that Mr R. E. Scott be elected to fill the vacancy on the 
Committee, on the retirement of Miss Helen Baker. This was seconded 
by Dr David Snow, and agreed by all. 


Any other business. [bis Editorial policy. Mr F. M. (Martin) 
Gauntlett had raised this subject, suggesting that the Club should make 
a representation to the Union that Jbis had become “‘unreadably boring 
and technically incomprehensible” to the average reader, and was 
primarily “‘aimed at professional biologists with a Ph.D. in statistics 
and mathematics’. He would like to see Ibis become more of a 
mainstream journal, with statistical tables and citations relegated to an 
appendix. 

‘The Chairman responded that it was not Club policy to comment on 
the editorial policy of the journals of others, and had invited a response 
from Dr Robert Prys-Jones, who had consulted with Dr Andy Gosler, 
Editor (designate) of [bis. 

Robert Prys-Jones read out a formal statement, dated 16 May, 
received from Andy Gosler, who welcomed the opportunity to outline 
his editorial vision for the future. He had taken note of these 
observations, and had some sympathy with the need to reduce the 


Meetings 133 Bull. B.O.C. 1998 118(3) 


increasingly obtrusive presentation of statistics in Jbis, but some 
statistical validation was necessary if /bis was to compete worldwide 
in the field of scientific ornithology and retain its position as one of the 
top three scientific journals, in a field of hundreds. A balance had to 
be struck between attracting the best scientific papers from the 
professional scientific community, and retaining the loyalty (and 
subscription income) of the average Membership. 

Dr Gosler said he had been involved with editing Jbis for only a few 
months and he had already in mind a number of editorial changes to 
make it more generally readable, but there was a backlog of papers in 
press, and it would be some time before these could be put into effect. 


No other items for discussion had been notified in accordance with 
Rule (12), and the meeting closed at 6.45 p.m. 


CLUB NOTES 


The eight hundred and seventy-fourth meeting of the Club was held on Tuesday, 17 
March 1998 at 6.15 p.m. 28 Members and 11 guests attended. 

Members present were: The Rev. T. W. GLADWIN (Chairman), Miss H. Baker, P. J. 
BELMAN, P. J. BuLL, D. R. CaLprer, Cdr M. B. CasEMENT rn, Prof. R. J. CHANDLER, 
S. J. FARNswortTH, A. Gipss, D. GRIFFIN, G. P. Jackson, J. A. JOBLING, R. H. KETTLE, 
I. T. Lewis, Dr J. F. Monk, D. J. Montier, Mrs A. M. Moore, R. G. Morcan, Mrs 
M. N. MULLER, R. C. Price, Dr R. P. Pr¥s-JoNgs, N. J. REpbMaAN, Dr R. C. SELF, P. J. 
SELLAR, N. H. F. Stone, C. W. R. Storey, A. R. Tanner, Dr D. R. WELLs. 

Guests attending were: Dr A. J. PRATER (Speaker), Mrs J. C. Buti, Mrs C. R. 
CASEMENT, Dr D. Foskett, Mrs J. M. GLapwin, Ms C. Horr, Mrs S. L. Lewis, Mrs M. 
MontTier, Ms A. Nussey, P. J. Moore, D. WITHRINGTON. 

On completion, Tony Prater presented an illustrated talk on ‘‘Waders’’, which was 
enthusiastically appreciated by all those present. 


The eight hundred and seventy-fifth meeting of the Club was held on Tuesday, 21 April 
1998 at 6.15 p.m. 36 Members and 20 guests attended. 

Members present were: The Rev. T. W. GLADWIN (Chairman), Miss H. BAKER, 
Captain M. K. BarritT rN, P. J. BELMan, I. R. BisHop, Mrs D. M. Brapb.ey, P. J. BULL, 
W.O.2 P. Carr RN, Cdr M. B. CASEMENT RN, S. E. CHAPMAN, Prof. R. A. CHEKE, G. S. 
Cow Les, N. J. Crocker, R. C. Dickey, A. Grass, D. GrirFin, G. P. Jackson, Dr J. B. 
KersLey, Dr C. F. Mann, Dr P. J. OLiver, L.A.(Met.), C. M. C. Patrick rn, Dr R. P. 
PrYs-Jones, N. J. RepmMan, S. J. Rumsey, R. E. Scott, Dr R. C. SELF, P. J. SELLAR, 
N. H. F. Stone, Cdr F. S. Warp rn, Prof. W. E. WaTErRs. 

Guests attending were: Ms G. BoNHaAM, Mrs S. Bourng, M. BraD.Ley, Mrs J. C. BULL, 
Mrs C. R. CaAsEMENT, Mrs P. A. CHAPMAN, Miss J. H. Cooper, Vice-Admiral Sir David 
Dosson, Mrs B. Grssps, Mrs J. M. GLapwin, Ms C. Horr, B. J. HuGuHes, A. J. MarTIN, 
Mrs M. Monrtier, P. J. Moore, P. Mutter, Cdr A. C. MurGatroyD Rn, B. O’BrIEN, 
Mrs J. Waters, M. WILSON. 

On completion, Dr Bourne gave an illustrated talk on ‘‘Birds and Islands’’. Bill first 
became interested in islands when evacuated to Bermuda in 1940-44, then during student 
expeditions to Rhum and the Cape Verde Islands in 1950 and 1951, and finally when sent 
to Cyprus on national service with the RAF, in 1956-58. Since islands often provide 
scope to make useful observations in a short time, he subsequently visited a variety, 
notably Gough and Chatham Islands, and Juan Fernandez. 

In general, islands fall into two classes—continental islands cut off from the main 
land-masses, with samples of their biota; and oceanic islands, mainly thrown up for a 
geologically limited period of a few million years by volcanic action at sea, and colonised 
by a less representative selection of organisms capable of crossing the water. The natural 
communities of islands in high latitudes were usually wiped clean by the glaciations, and 


Meetings 134 Bull. B.O.C. 1998 118(3) 


there has been inadequate time in the 10,000 years since the last of them to show much 
variation. But those of islands in lower latitudes have often been able to respond to 
changes in climate by moving up and down hill, so that they often support ancient relicts, 
which have achieved such a high level of evolution that their nearest relatives are now 
doubtful. 

Thus, while the northern Atlantic islands hold about 60 endemic forms, half in Britain 
and Ireland, they mainly show the extreme development of general geographical trends of 
variation and colour, and it seems doubtful if any really deserve recognition as endemic 
species. On the other hand, while the continental Mediterranean islands only hold about 
a third as many races, possibly because they are insufficiently isolated (whereas they used 
to hold many more endemic mammals), they do hold three highly distinct species (13%): 
the Corsican Nuthatch Sitta whiteheadi, Cyprus Wheatear Oenanthe cypriaca and Cyprus 
Warbler Sylvia melanothorax. The first is adapted for montane forest, and the other two 
for an extreme Mediterranean climate. 

Compared with this meagre showing for the NW Palearctic continental islands, there is 
a remarkable concentration of endemic forms, apparently often of ancient and diverse 
origin, in the west African oceanic islands. These include, at a conservative estimate, 
some 48 endemic forms, of which eight (17%) rate as species, in the Canaries and 
Madeira, and 18, of which four (25%) rate as species in the Cape Verde Islands. The 
species moreover include some truly remarkable birds, such as the two Laurel Pigeons 
Columba junoniae and C. trocaz, which might be aberrant derivatives of the Woodpigeon 
C. palumbus, which have undergone further local evolution, or be left over from laurel 
forests in the Mediterranean area during the tertiary, and also early derivatives of 
ancestral swifts, larks, pipits, chats, warblers and sparrows. 

‘The situation in the tropical archipelagoes becomes too complicated to summarise 
easily, and to study insular variation in its extreme form it is necessary to go further south 
to the more isolated sub-antarctic islands. Here, for example, all the seven landbirds of 
the Tristan—Gough group show such marked local evolution that they were formerly 
placed in five endemic genera of uncertain affinities thought to represent ancient relicts. It 
now seems obvious that the two gallinules Porphyriornis nesiotis and P. comeri are actually 
just large flightless Moorhens Gallinula chloropus, however, while the smaller flightless 
Inaccessible Rail Atlantisia rogersi sometimes has distinctive spots like those of the South 
American Spot-winged Crake Porzana spiloptera. 

Similarly, the single Gough passerine Rowettia goughensis appears to be a South 
American finch Melanodera masquerading as a thrush, whereas on Tristan and its 
outliers, where there are three passerines, there is a thrush Nesocichla eremita which may 
be a South American Groundscraper Turdus litsitsirupa adapted to suck eggs, and large 
and small canaries Nesospiza acunhae and N. wilkinsi, both derived from small South 
American yellow-finches of the genus Sicalis. 

There is a need for more study of these remarkable and often threatened birds, and it 
might be helpful if more attention could be devoted to their relationships and the reasons 
why they have come to differ from their allies, instead of unreal hypotheses about their 
phylology and island biogeography. 


The eight hundred and seventy-sixth meeting of the Club was held, following the Annual 
General Meeting, on Tuesday, 19 May 1998 at 6.45 p.m. 23 Members and 7 guests 
attended. 

Members present were: The Rev. T. W. GLADWIN (Chairman), Miss H. Baker, I. R. 
BisHop, Cdr M. B. CASEMENT RN, Prof. R. J. CHANDLER, D. J. FisHER, F. M. 
GAUNTLETT, D. GriFFIN, J. A. JoBLING, K. J. KazmrerczaKk, M. B. LANCASTER, Dr 
J. A. K. Metprum, D. J. Montiger, Mrs A. M. Moore, Mrs M. N. Mutter, M. L. 
PaLInG, Dr R. P. PrY¥s-Jongs, N. J. RepmMan, R. E. Scott, P. J. SELLAR, Dr D. W. 
Snow, N. H. F. Stone, C. W. R. STOREY. 

Guests attending were: Mrs G. BonHAM, Mrs M. H. Gaunt ett, Miss J. H. Cooper, 
Mrs J. M. Giapwin, Mrs M. Montier, P. J. Moore, Mrs P. Pon. 

After dinner, Members gave a series of short talks on subjects of topical interest; the 
following is a brief synopsis. 


“Hybrid Lapwings’’. Prof. Richard Chandler showed photographs of two different 
apparent hybrids between Blacksmith Plover Vanellus armatus and Spur-winged Plover 
V. spinosus. The two species are regarded by most authorities as being closely related. 


Meetings 135 Bull. B.O.C. 1998 118(3) 


Both individuals were seen at Lake Nakuru National Park, Kenya, in an area where the 
two species are sympatic, one by the speaker in 1990, the other by Dave Richards in 1997. 
Both appeared to be adults, and showed plumage patterns intermediate between the 
presumed parents, with buff-grey upperparts and wing coverts, black on the head 
(distributed differently in the two individuals), white hind-neck collar with black lower 
margin, and white underparts with a black band across the belly. Both had black bills; the 
first individual had black legs, the second grey. 


‘Duetting in the Gambia’’. Mr P. J. Sellar illustrated with slides and tape recordings 
the recent work undertaken by a St Andrews University field project in The Gambia to 
determine whether the lead songs in duetting are performed by the male or female in the 
Oriole Warbler Hypergerus atriceps and the Barbary Shrike Laniarius barbarus. In both 
cases the male was found to lead the duet. 


Mr M. B. Lancaster showed a series of slides to illustrate the coloration of Plain Swift 
Apus unicolor seen in Tenerife during almost annual visits there since 1989. He had 
never knowingly seen Common Swift A. apus there, but the presence of several swifts 
with well marked throat patches had prompted closer attention, and he had taken a series 
of photographs. Plain Swifts had always appeared to the speaker as ‘brown’, especially 
towards dusk and dawn, and when viewed from below, whereas Common Swift is ‘black’. 
Pallid Swift A. pallidus had been identified without difficulty—much paler with a large 
throat patch. T. Clarke (resident in Tenerife) had commented that Plain Swift may also 
show a pale throat, and that differentiation of the three Apus spp. was not easy. 
Comments from the audience suggested that slight over-exposure may have been the 
main contributory factor to the brown effect, while the pale areas could have been 
reflected light. The speaker commented that photographs of White-throated Swift 
Aeronautes saxitalis, taken in Canada under similar lighting conditions, and using the 
same film, produced a ‘grey’ image. 


“Cyanopica—a mystery solved’’. Joanne Cooper said that the origin of the disjunct 
distribution of Azure-Winged Magpie Cyanopica cyanus has never been fully explained. 
The species is found in eastern Asia, across China, Japan and Korea, and in western 
Europe, in the Iberian Peninsula. T'wo main explanations have emerged to account for 
this pattern: firstly, that it developed during the last glacial period, as parts of a once 
continuous population were isolated in refugia and intervening populations became 
extinct; secondly, that the species was introduced into Iberia during the 16th Century by 
Portuguese sailors returning from the Far East. The recent identification of fossil remains 
of Cyanopica cyanus from Neanderthal occupied sites in Gibraltar reveal its presence in 
Iberia during the last glacial period. This evidence provides proof that the species was not 
introduced into Iberia in recent times, but is instead a refuge relict. 


“El Nino”. Mr R. E. (Bob) Scott reminded members that this is a phenomenon of 
extreme weather conditions that occurs over an ill-defined 5—7 year cycle. The effect is a 
rise in seawater surface temperature that can cause dramatic environment changes, of 
very variable intensity. He had visited three distinct geographical areas during the 
autumn/winter of 1997/98 and recorded ‘abnormal’ weather patterns, which had had a 
major effect on the numbers and bird species seen there. 

In the Seychelles, Indian Ocean, the rains had started unusually early, in August, 
and the rains were still falling when he visited in October 1997. The heavy rainfall had 
caused serious damage to property, roads and loss of life. The rains do not normally 
start until November or December. On the Gulf coast of Texas, U.S.A., in November 
1997, the weather was unseasonably cold, with snowfalls well south in the State. 
Strong, cold winds on the coast contrasted with the balmy weather normally to be 
expected at that time of the year. Finally, on the Caribbean coast of Costa Rica, 
Central America, in January 1998, the season was unusually dry with normally 
navigable rivers barely passable by boat. In all three examples local residents credited 
these extremes to E] Nino. 

Members of the audience commented that several localities visited recently seemed to 
be recording their ‘wettest’, ‘coldest’ or ‘driest’ seasons for several years. 


Joint BOU/BOC/NHM/BirdLife Conference, 12-14 November 1999. Dr R. P. 
(Robert) Prys-Jones announced brief details of a two-day conference on the subject 
“Why Museums Matter; Avian Archives in the Age of Extinction’’, planned to take place 


Meetings 136 Bull. B.O.C. 1998 118(3) 


at Green College, Aston Clinton, less than 3 miles from the NHM Bird Group, Tring. 
The aims of the conference will be to highlight the continuing and changing importance 
of museum specimens, and associated library resources, vocalisations etc., to bird 
research and conservation; and to discuss ways in which the information contained within 
this resource can best be made available to a wider public. It will comprise an array of 
invited talks intended to cover most of the main aspects of this museum resource, 
together with some offered talks which must relate closely to the conference them. 

Directly following the conference, from the Sunday evening until the afternoon of 
Monday 15 November, there will be a one-day workshop at the Tring Museum site, open 
only to curators and managers of bird collections, on the subject of “Increased 
Co-operation between Museums, especially in Europe’’. It is very much hoped that 
representatives from most major bird collections will wish to attend both the workshop 
and conference, but space will be quite limited. A set of proceedings, comprising both the 
conference papers and a summary of the workshop discussions,:is planned for the BOC 
Occasional Papers series, during the year 2000. 


The eight hundred and seventy-seventh meeting of the Club was held on Tuesday, 14 
July 1998 at 6.15 p.m. 26 Members and 14 guests attended. 

Members present were: The Rev. T. W. GLADWIN (Chairman), Miss H. Baker, P. J. 
BELMAN, I. R. BisHop, Mrs D. M. Brapiey, D. R. CaLpEr, Dr M. Carswe.., Cdr M. B. 
CASEMENT RN, S. J. FARNSWORTH, D. J. FisHER, A. Gipsps, D. GRIFFIN, G. P. JACKSON, 
J. A. JoBpLinc, R. H. Ketrie, M. B. LANcaAster, D. J. MonTiER, R. G. Morcan, Mrs 
M. N. Muuier, R. E. F. Peat, Dr R. P. Prys-Jonges, N. J. REpMaNn, R. E. Scott 
(Speaker), Dr R. C. SELF, P. J. SELLaR, C. W. R. STOREY. 

Guests attending were: Ms G BonHam, Mr M. J. BrapLey, Mrs J. B. CALDER, Mrs B. 
Gipss, Mr R. GiLtBey, Mrs J. M. GLtapwin, Ms C. Horr, Mrs M. Monrtier, Mr C. A. 
Mutter, Mr B. O’Brien, Mr A. R. E. PEAL, Prof. D. E. Pomeroy, Mr and Mrs B. SacEe. 

After dinner, Mr R. E. Scott entertained Members with a lively talk, which amply 
fulfilled his title ““Bird reminiscences in a lighter vein’’. It was filled with amusing 
personal anecdotes and his readings of extracts, including poems, from a wide range of 
sources. 

Bob opened by recalling how his first knowledge of the countryside was when he was 
evacuated from London to Wiltshire, in the 1940s, and his lifelong interest in birds was 
triggered by his introduction to a tame Jackdaw. Treasured as his first bird books were 
the identification guides of F. B. Kirkman, James Fisher, T. A. Coward, Richard Fitter, 
and Peterson’s Field Guide, which transformed birdwatching throughout Europe in the 
1950s. The poem in Coward’s Vol. 3 Flying Machines by the Little Stint was responsible 
for his first interest in bird migration. 

His early work at the British Museum (Nat. Hist.) involved more time in the Bird 
Room than in Platyhelminths where he was employed! He abandoned this to work for the 
RSPB at their Dungeness Reserve and neighbouring Bird Observatory. There he became 
involved in studies sponsored by the Central Electricity Generating Board (CEGB) in 
support of conservation issues threatened by the construction of Dungeness A nuclear 
power station. That public enquiry lasted a mere one and a half days. He also took an 
active part assessing the mortality caused by power lines, and the effects on seabirds of 
the Torrey Canyon oil spillage disaster. Bob’s reminiscences were spiced with hilarious 
accounts of the well-meaning, but disastrous, attempts by members of the public to assist 
with this work. 

He subsequently moved to Sandy to become head of RSPB Reserves Management, 
where the predator/prey relationships of birds were a constant source of conflict between 
vested interests. In his retirement, he now travels widely as a leader and guide of bird 
tours, where bizarre and humorous situations continue to provide material for 
after-dinner talks. 


Accounts £37 Bull. B.O.C. 1998 118(3) 


British Ornithologists’ Club 
Financial statements for the year ended 31 December 1997 


Trustees’ Responsibilities 


Under the Charities Act 1993, the Trustees are required to prepare a 
statement of accounts for each financial year which give a true and fair 
view of the state of affairs of the charity at the end of the financial year 
and of the incoming resources and application of resources in the year. 
In preparing the statement the Trustees are required to: 


@ select suitable accounting policies and then apply them consist- 
ently; 

@ make judgements and estimates that are reasonable and prudent; 

@ state whether applicable accounting standards and statements of 
recommended practice have been followed, subject to any material 
departures disclosed and explained in the statement of accounts; 

@ prepare the financial accounts on the going concern basis unless it 
is inappropriate to presume that the charity will continue its 
operations. 


The Trustees are responsible for keeping proper accounting records 
which disclose with reasonable accuracy at any time the financial 
position of the charity and to enable them to ensure that any statement 
of account prepared by them complies with the regulations under 
section 41(1) of the Charities Act 1993. They are also responsible for 
safeguarding the assets of the Trust and hence for taking reasonable 
steps for the prevention and detection of fraud and other irregularities. 


BRITISH ORNITHOLOGISTS’ CLUB 
Registered Charity No. 279583 
Balance Sheet as at 31 December 1997 


1997 1996 
Notes if fi if £ 
Tangible Fixed Assets ........................ 2 40 50 
Investments 
PM MIAI ERE ETIRANTES 2-9 5250-5 fc. ccoccscceustusecce ces 3 202,878 173,446 
Current Assets 
Stock of publications........................4- 100 100 
Cash at bank and in hand................... 5,099 4,660 
Se or et 56,536 56,788 
JOLT SCT ET UT |. Si a 480 60 
Lf oh oe eas 2,462 81 
64,677 61,689 
Current Liabilities 
Subscriptions in advance...........2..00..0. (5,437) (4,740) 
Creditors falling due within one year.. (3,985) (4,673) 
55,255 52,276 


TOTAL ASSETS £258,173 E22 a TTL 


Accounts 138 Bull. B.O.C. 1998 118(3) 


Funds 
Winrestricte dress: tsconpecveysesssseehecsouseeseete cs 4 Dallas 219,308 
FRESEPICEC Foo 55s eile ose cence yaaaue Me catunmemae tease 5 7,0504 51; 6,464 


£258,173 £225,772 


Approved and signed on behalf of the Trustees 
T. W. GLADWIN 
Chairman 


21 April 1998 


Statement of Financial Activities Year Ended 31 December 1997 


Unrestricted Restricted 


Funds Funds Total Total 
1997 1997 1997 1996 
Notes fe if vb £ 
INCOME 
Subscriptions 
Memb ers) Sh 3 ited dtisice «Acie tos Soe eEe ack th ae eS S eRe ae 6,559 — 6,559 4,961 
Non-member subscribers..............cccceccecseeeccseeeeveees BBS) —_— 3,89 SSIS} 
Income tax recoverable under Deeds of Covenant 328 — 328 308 
10,022 — 10,022 8,582 
Donations}: 212s Rep eS a oeet + bod. 50 165 215 385 
Sponsorship received ..................ccccccseeeceeeeeneeeeneeeees — —_— —— 2,488 
Investment iINCOME............... 0... eeccecceccesecseceaseaeceaees 
Herbert Stevens Trust Fung .............:cccsesssecesseeeeees 10,167 — 10,167 8,334 
Income tax recovered in respect of prior year ......... 1,426 — 1,426 1,578 
11,593 11,593 9,912 
Barrington Trust Fund COIF Income Shares......... 39 — 39 38 
Interest FeCeivieds.cuscscckensstwcheaaagecneee nesses Rese eee 3,386 421 3,807 3,146 
15,018 421 15,439 13,096 
Sales of Publications 
Bulletintsetast hl edhe eee dd. lace Re hee 1179 —_ 1,179 2,114 
Otherpubblicationss. Haye dew ee. ao eee ene eee ee ee 3,010 — 3,010 1,458 
4,189 — 4,189 3,572 
Meetings 3,726 — 3,726 3,131 
Other income 
Club»tresyandibrooches sare. rig-ckes pase eseeen eres te eeeen-iote 107 —_ 107 Ue 
Miiscellameous: x -prcttheds seat iacp Saicee Meee eaten ates eh ee cance — — = 13 
TOTALAINCOMB see ees See aR eae 33,112 586 33,698 31,344 
EXPENDITURE 
Direct charitable expenditure 
Meetings 
Room hire, speakers’ expenses, etC...........0.....:0eeeeees 1,226 — 1,226 203 
BOC Bulletin 
Production, printing and distribution 14,877 — 14,877 13,178 
Other publications 
Production and publication.............cccccseceeeeneeeeeenees 7,904 — 7,904 972 
Buture! publicatiomsmnccscstecse specs eee eas eee ee eee 500 — 500 —_ 
Publicity, postage and packing..................2eeeseeseeeee 358 —_ 358 40 
Other expenditure 
Meetings 
Restaurant. 2h eee eet oA eran eM Ren roel 3,519 3,519 320M 
Administrationyrcoscessesss ase yee eee reece esate 6 2,345 2,345 4,607 


MOMATE: EEXCPIE IND TU RIBy 2 oereeuesencre seein reece 30,729 = 30,729 22,251 


Accounts 139 Bull. B.O.C. 1998 118(3) 


EXCESS OF INCOME OVER EXPENDITURE... 2,383 586 2,969 9,093 

Appreciation in value of investments...................08..055 29,432 — 29,432 9,845 
31,815 586 32,401 18,938 

TOTAL FUNDS brought forward at 1 January 

ea AUR no) Sec Baen eee ee Sadat an Stead saa wez ecko ssn 219,308 6,464 225 te: 206,601 

Prior year adjustment to Barrington Trust Fund........ — — — 233 

TOTAL FUNDS at 31 December 1997 ...................... EPS £7,050 £258,173 £225,772 


NOTES TO THE ACCOUNTS 

YEAR ENDED 31 DECEMBER 1997 

1. ACCOUNTING POLICIES 

(a) Basis of Accounts 
The financial statements are prepared under the historical cost convention. 

(b) Investments 
Investments in the Herbert Stevens and Barrington Trust Funds are shown in the Balance Sheet at market 
valuation. This represents a change of policy in the case of the Barrington Trust Fund Income Shares 
previously stated at cost, being £577. The cumulative appreciation in value up to 1 January 1996 of £233 and 
for 1996 of £61 have been treated as prior year adjustments and comparative figures amended accordingly. 

(c) Depreciation 

Depreciation is calculated to write off fixed assets over their expected useful lives at an annual rate of 10% on 

cost. 

Publications 

The cost of publications is written off in the Income and Expenditure Accouni as incurred except for a nominal 

stock value of £100 carried in the Balance Sheet. 


(d 


— 


Projection 
equipment 
2. TANGIBLE FIXED ASSETS 
RaiseeEnneaaiary and ot Mecember L997. ......:-..ccescensccesacesceeccetanseionsiow 100 
Accamulated depreciation at 1: January 1997........2...ecckece..cceeeoccceeosseeees 50 
SEAGER PDP RE (Gor Eyes eee cee ce enna EES coc ce ene eee 10 
Pee Pat OO) see SE AE SEE Jk RR SS ee 60 
nen uorcararmiceteyn wecember 1997... 5......:5.c.2.--08. eonedectcsecssesessesssseek! £40 
Neibookwwiluc at sieWecember 1996 «.......c.c-2-..0....00.e-cosoecccesseveseseoaedecss £50 
1997 1996 
3. INVESTMENTS—at market value rh if 
SCEPD ES! SPRUE PSM io isjel 8 rie tc | eee ee ee en eee eee er Pee ee 201,854 172,575 
S-PF o rpini 7 TS o EO ra ee ee oer eee ee 1,024 871 
£202,878 £173,446 
General Herbert Stevens Barrington 
fund Trust Fund Trust Fund Total 
4. UNRESTRICTED FUNDS if £ if £ 
Balance at 1 January 1997.................. 45,862 172,575 577 219,014 
Prior year increase in investment to 
market value (Note 1b) ...................... as — 294 294 
45,862 172,575 871 219,308 
Appreciation in value of investments 
LES o VE ee eee — 29,279 153 29,432 
Surplus of income over expenditure... 2,383 — — 2,383 
£48,245 £201,854 £1,024 yey bs (A283 
Publications Bird Atlas of 
Fund Uganda Fund Total 
5. RESTRICTED FUNDS £ eR iff 
Balance aed jJaritiary (997 0 oF. s.3.-ccccsen-cse. 3,867 2,597 6,464 
Womttaas . .123.. FOE. 1S. Se Ws..t: 165 —_ 165 
CPL SST ee ps 5 ae Se oe 253 168 421 


Errata 140 Bull. B.O.C. 1998 118(3) 


1997 1996 
6. ADMINISTRATION EXPENSES include: £ 
Auditeand! Independentiexammiationl fees -.-n-erssser eee teaneenssase-ce eer teee eee tee 550 1,059 
Depreciationvol tangible fixe dlasseiseresteae- tee eee neee eee ee eee eee 10 10 


7. REIMBURSEMENT OF EXPENSES 
Committee members are reimbursed for expenses incurred by them on behalf of the Club. The amount reimbursed 
during the year was £1,271 (1996 £1,112). 


INDEPENDENT EXAMINERS’ REPORT TO THE TRUSTEES OF THE 
BRITISH ORNITHOLOGISTS’ CLUB 


This is a report in respect of an examination carried out on the accounts set out on pages 137 to 140 under Section 43 
of the Charities Act 1993 and in accordance with directions given by the Charity Commissioners under Sub-section 
7(b) of that Section. 


RESPECTIVE RESPONSIBILITIES OF TRUSTEES AND INDEPENDENT EXAMINERS 

As described on page 137 the Trustees are responsible for the preparation of the accounts and they consider that 
Charities Act 1993 Section 43(2) (audit requirement) does not apply. It is our responsibility to carry out procedures 
designed to enable us to report our opinion. 


BASIS OF OPINION 

This report is in respect of an examination carried out under Section 43 of the Charities Act 1993 and in accordance 
with the directions given by the Charity Commissioners under Section 43(7)(b). An examination includes a review of 
the accounting records kept by the Charity Trustees and a comparison of the accounts presented with those records. It 
also includes a review of the accounts and making such enquiries as are necessary for the purposes of this report. The 
procedures undertaken do not constitute an audit. 


OPINION 

No matter has come to our attention in connection with our examination which gives us reasonable cause not to believe 
that in any material respect: 

(i) accounting records have been kept in accordance with Section 41 of the Charities Act 1993; or 

(ii) the accounts accord with those records; or 

(111) the statement of accounts complies with the accounting requirements of the Charities Act 1993. 

No matter has come to our attention in connection with our examination to which, in our opinion, attention should be 
drawn to enable a proper understanding of these accounts. 


Sevenoaks PORRITT RAINEY & CO. 
Kent Registered Auditors 
TN13 1XR Chartered Accountants 


30 April 1998 


Errata J 


Bull. Brit. Orn. Club 117(1): 77-79 (1997). Mariano Ordano and Andrea Bosisio have 
pointed out the following error in their paper Historical records of threatened and near 
threatened Argentinian birds from Museo Provincial de Ciencias Naturales “Florentino 
Ameghino”’ of Santa Fe, Argentina: “Specimens from Museo Provincial de Ciencias 
Naturales ‘‘Florentino Ameghino”’ collection’s classified as Coryphaspiza melanotis 
(Black-Masked Finch), published in Bull. B.O.C. 117(1): 77-79, are Donacospiza 
albifrons (Long-Tailed Reed Finch), whose numbers are MFA-ZV-1670, 1359, 639 and 
638. We acknowledge the mistake in the specimen’s identification and apologise for the 


> 


error. 


Bull. Brit. Orn. Club 118(2), Plate 1. Due to a printing error, the caption was omitted 
from Plate 1, which accompanied the report of Ian Burrows’ talk to the 873rd meeting of 
the Club. This should read “Feline Owlet-nightjar Aegotheles insignis, Mount Robinson 
1,600 m, Western Province, Papua New Guinea, 24 July 1994 (lan Burrows)’. 


Bull. Brit. Orn. Club 118(2): 67-82. In Frank Lambert’s paper A new species of 
Amaurornis from the Talaud Islands, Indonesia, and a review of taxonomy of bush hens 
occurring from the Philippines to Australasia, on line 12 under “Diagnosis” (p. 74), “A. 
moluccanus”’ should be replaced with “‘A. magnirostris’’. 


Plate 3. Top left—Otus alius, sp. nov., compared with the related taxa which it most 
closely resembles in plumage (all probably illustrated for the first time): top 
right—grey-morph O. magicus tempestatis (Wetar Island); bottom left—O. [magicus| 
kalidupae (‘Tukangbesi Islands); bottom right—O. [m.] swlaensis (Sula Islands). All other 
taxa are more dissimilar in aspect and hence are not illustrated. Irides of alius are 
assumed to be yellow based on iris colour of most related taxa. Original painting by Larry 


B. McQueen. 


P.C. Rasmussen 141 Bull. B.O.C. 1998 118(3) 
A new Scops-owl from Great Nicobar Island 


by Pamela C. Rasmussen 
Received 6 Fuly 1998 


A single red-morph Scops-owl collected on Camorta, central Nicobar 
Islands in 1873 formed the basis for the 1876 description: of Ephzaltes 
nicobaricus Hume, and a second, nearly uniform rufous specimen was 
obtained from the same island in 1874 (Gurney 1884). The taxon was 
treated as a racial synonym of Otus sunia modestus or O. s. malayanus by 
some, and by others as a subspecies of Stresemann’s (1925, 1939) 
extremely varied, nearly worldwide O. scops. No further Scops-owls 
were reported from the Nicobars until one was collected on Great 
Nicobar in 1966 (see Plate 3, upper left). Ali & Ripley (1969: 265) 
footnoted this specimen as O. s. nicobaricus, observing that it was 
*... considerably larger than modestus ... with which Peters (1940) 
would synonymize it.’ Abdulali (1972: 106) noted it ‘does not agree 
with the original description ... The wing lacks the white edge 
common to the other races and the lower belly is also as profusely 
marked as the breast ... Though required for all races of O. scops ... 
the first primary is not longer than the eighth’. Its long wings led 
Marshall (1972) to believe the 1966 specimen was not sunza, and after a 
second specimen was collected on Great Nicobar in 1977, Abdulali 
(1978: 759) stated ‘Marshall who has been studying this in detail is sure 
that it is an undescribed form but cannot decide of what species’. 

Marshall's (1978) discovery that dissimilar taxa from widely disjunct 
islands had similar calls led him to postulate that Otus magicus 
(formerly restricted to Wallacea) is highly polytypic, with ‘...a history 
of long flights over oceans and recent colonizations’. He tentatively 
placed the Great Nicobar bird in an expanded Otus magicus, citing ‘.. . 
morphological similarity to zmsularis ...’ of the Seychelles. He stated 
that the unnamed taxon lacks eartufts, provided a brief partial 
description (but no name), and included it with ‘... taxa of unknown 
song ... hence of unknown affinities .. . listed with a guess as to their 
possible allocation ...’ (Marshall 1978: 17, 32). It has subsequently 
been treated either as O. s. nicobaricus (Ali & Ripley 1981, Ripley 1981) 
or O. magicus (Marshall & King 1988), until it was recently listed as 
‘Nicobar Scops-Owl Otus sp.’ (King 1997: 26). 

In late 1996, I examined both Great Nicobar specimens, now at the 
Bombay Natural History Society (BNHS), and that same week both 
Camorta specimens at The Natural History Museum, Tring, U.K. 
(BMNH), realizing then that specimens from these two islands were 
very different. Subsequent comparison with all specimens of each taxon 
in the manadensis superspecies (Marshall 1978) in the museums listed 
in the Acknowledgements has led to confirmation that the unnamed 
Otus is distinct morphologically in a number of characters. Given this, 
its isolation (Fig. 1), and an emerging pattern of high geographic 
differentiation among island Otus, the Great Nicobar bird cannot be 


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Bull. B.O.C. 1998 118(3) 


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P. C. Rasmussen 143 Bull. B.O.C. 1998 118(3) 


specifically aligned with magicus or any other taxon, but requires 
recognition as a distinct species. 


NICOBAR SCOPS OWL Otus alius, sp. nov. 


Holotype. Bombay Natural History Society No. 22578, adult male (see 
Plate 3, upper left) from Campbell Bay, Great Nicobar Island, 7°00'N, 
93°50’E, India; collected 3 March 1966 by Humayun Abdulali. 

Diagnosis. A medium-sized Otus with predominantly warm brown, 
mostly unstreaked and finely barred plumage; ear tufts evenly and 
finely barred, rounded, and of medium length; facial disk indistinctly 
paler than rest of plumage, lightly barred at its lower edge; dark border 
to facial disk not prominent; bill medium-sized and mostly brownish; 
white scapular spots rounded, nearly restricted to outer web, with 
broad black tips; most distal portion and much of rear edge of tarsus 
bare; toes and claws relatively large. 

Otus alius is closest in morphology to some members of the 
manadensis superspecies of Marshall (1978), from all of which it differs 
in possessing scapular spots that are bordered above and below by 
broad black bands, giving a rounded shape to the white centres. It 
differs additionally from adults of all Asian taxa in possessing a 
combination of finely barred overall plumage, entirely lacking streaks 
above, and having a marked reduction of ventral streaking, over which 
the heavy tricoloured barring predominates; and in having relatively 
large claws. Although sample size is insufficient for statistical testing, 
its wing formula differs from all taxa Marshall (1978) placed in magicus 
(but not O. enganensis or O. umbra) in that P3 (from outermost) is 
longer, while the inner secondaries are shorter (‘Table 1), and like the 
latter two only, the bill is mostly brownish. Additional differences are 
given in the ‘Remarks’ section. 

Distribution. Both specimens were collected near sea level at 
Campbell Bay, Great Nicobar Island. There are no other published 
reports of Scops-owls from Great Nicobar. 

Description of the holotype. Capitalized colours are from Smithe 
(1975); colour comparisons were made under weak fluorescent light, 
with diffused sunlight from an open window. 

Sides of forehead above supercilium finely barred Cinnamon (39), 
whitish, and Burnt Umber (22); base of central crown feathers Clay 
Color (26), central streaks Fuscous (21); base of poorly-marked whitish 
supercilium feathers paler than Cinnamon, with narrow, widely-spaced 
Burnt Umber bars; base of ear tufts Cinnamon, narrowly barred Dark 
Grayish Brown (20); ear tufts rather short, prominent, with broad 
feather tips; bases of facial feathers at sides of bill more rufescent than 
Cream Color (54); bases of rictal bristles Buff-Yellow (53), at least tips 
blackish; short feathers below and in front of eye vaguely dark-barred 
with pale Cream Color bases; facial disk behind eye Cinnamon with 
whitish shafts and dusky mottling, rear and lower portion whitish with 
fine Dark Grayish Brown barring and mottling, narrow, indistinct disk 
rim Dark Grayish Brown on lower and rear edges; only slight 
extensions to rear auriculars despite fresh plumage; chin feathers 


P. C. Rasmussen 144 Bull. B.O.C. 1998 118(3) 


coloured as feathers at bill sides; throat pale buff with fine dark barring 
near tips of longer chin feathers. Rest of crown, nape, and hindneck like 
central crown, but with streaking becoming obsolete and narrow dark 
bars prominent on rear crown. 

Mantle through lower back close to Raw Umber (23) in aspect, dark 
bars ca. 2 mm wide, slightly paler than Fuscous, light bars ca. 1.7 mm 
wide, mostly Cinnamon but paler near bases. Scapular spots rounded, 
mainly white encircled by Dark Grayish Brown (blackish) bands, and 
surrounding base colour Amber (36), with fine Burnt Umber markings, 
scapulars Cinnamon near bases. Tertials have broad (ca. 4.1 mm) 
indistinct bands between Fuscous and Olive-Brown (28) and narrow 
(ca. 1.3 mm) Smoke Gray (45) bands; largest feathers of alula with bars 
ca. 2.5 mm, Cinnamon and paler, medium bars ca. 4.0 mm and stippled 
darker, narrow dark bars Fuscous; outer primary (P1) has lightest 
bands slightly paler than Cinnamon, medium bands slightly paler than 
Olive-Brown, and darkest bands Fuscous; light bands paler on inner 
primaries. Uppertail coverts closest to Raw Umber (23), with fine dark 
bars of Burnt Umber giving vermiculated aspect, uppertail surface with 
palest bands of Cinnamon, medium bands finely speckled with dark, 
giving Olive-Brown aspect, darkest bars Fuscous; banding pattern 
fairly strong on uppersides of rectrices. 

Upper breast Cinnamon with ca. 1.5mm distinct Fuscous bars 
distally, ca. 1.2 mm Fuscous shaft streaks; lower breast Cinnamon with 
narrow bars between Olive-Brown and Fuscous, on breast sides shaft 
streaks are Fuscous, a few as wide as 2.2 mm, some bars slightly paler 
than Cinnamon; centre of belly with more white, white bands ca. 4 mm 
broad, Cinnamon bands ca. 3 mm, blackish bars ca. 0.7 mm, narrow 
white shaft streaks; tips of bars on lower flanks deteriorate into 
vermiculations, otherwise they are broad Cinnamon bands with narrow 
white and fine blackish bars. Undertail coverts rufescent whitish for 
most of their length but with tips barred as is belly, although rufous 
bars are closer together. Tarsal feathers mostly Cinnamon with fine 
Fuscous bars; bare patch on upper hind tarsus, with a narrow bare strip 
along the upper third, feathering meeting ventrally on medial section; 
dorsally, feathering ends 8.6 mm short of joint formed by phalanges 
1-2 of third digit. 

Dried cere dull yellowish-brown; maxilla tip and distal half cutting 
edge blackish-brown, most of sides of maxilla dull yellowish-brown, 
base and culmen ridge paler and yellower; mandible dull yellow with 
cutting edge and base dark brown. Dried toes dark yellowish-brown, 
claws mostly dusky horn, black distally, grading near bases to paler 
yellowish-brown. Narrow ring of bare orbital skin blackish. No data on 
soft part colours of fresh specimens or living birds. 

Description of the paratype. The female paratype is similar overall to 
the holotype, but is slightly larger, more boldly marked, with more 
white on lower underparts. Dark bands on forehead broader, facial disk 
slightly darker and more heavily marked behind eye; longest throat 
feathers with coarser and more widely spaced blackish barrings; mantle 
more coarsely barred, with larger, whiter spots basally; alula more 
contrasting; upper breast more strongly marked and lower breast more 


Bull. B.O.C. 1998 118(3) 


145 


P.C. Rasmussen 


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P. C. Rasmussen 147 Bull. B.O.C. 1998 118(3) 


variegated; flanks have white bands broadest and blackish bands 
heavier; upper tail more coarsely speckled, less banded; primaries with 
pale bands lighter; tarsal feathering paler and more speckled. Both 
types resemble each other more closely than either does any specimen 
examined of other taxa. 

Measurements of the holotype (mm). Wing, flattened and straightened 
(161.0); central rectrix (74.4); tarsus (28.6); culmen from skull (21.7); 
culmen, from anterior edge of cere (12.8). For additional measurements 
and those of related species, see T‘able 1; sample sizes are insufficient 
for statistical testing. 

Specimens. Great Nicobar: BNHS (holotype); BNHS 24411 (9; 
paratype) from type locality (formerly Zoological Survey of India [ZSI] 
No. 33171, collected 2 April 1977-by S. S. Saha for K. K: Tiwari). 

Etymology. ‘The name alius, which is Latin for ‘other’ (this being 
another Scops-owl from the Nicobar Islands), encapsulates the family 
name of Mr Humayun Abdulali, who first collected this species, and 
contributed a great deal to Indian ornithology, and in particular that of 
the Andaman and Nicobar islands. The common name ‘Nicobar 
Scops-Owl’ is appropriate and already in use solely for this taxon (King 
1997: 26). 


Remarks 
Additional differences 


From O. beccarii (n=2 specimens examined and measured, and 
photographs of type; Biak I.), alms differs greatly by its less 
contrasting, paler, browner, less fluffy plumage; mantle more definitely 
barred; more slender bill; streaked forecrown; more definite eartufts; 
less distinct facial disk; somewhat streaked below; primary coverts and 
alula not dark and unmarked; and shorter, more prominently banded 
tail. The highly isolated Otws beccarii is so distinct in plumage that even 
Stresemann (1925) and Mayr & Meyer de Schauensee (1939) gave it 
specific status, while Marshall (1978) lumped it in magicus without 
examining specimens until his manuscript was in proof. 

From Otus insularis (Seychelles, n=6; colour plate of type, Tristram 
1880), alius differs markedly in its much smaller size, especially skull 
and bill (Table 1); less prominent dark eye patch; eartufts not mainly 
buffy and blotched; less rufflike head feathers; less prominent facial 
disk edge; lack of buff-spotted hindcollar; scapular spots not narrowly 
black-edged and lined; relatively longer wings; much shorter, slighter, 
more extensively feathered tarsi; and relatively longer tail with broader 
central rectrices (Table 1). Otus insularis diverges most from alius of 
any of the taxa considered here in its very heavy streaking below and 
heavy blotching above. All adult specimens and photos of three live 
birds are similar, but differ from other ‘magicus’ taxa, and insularis 
is separated by so great a distance (Fig. 1) that others regard it as dis- 
tinct (Watson 1980, Collar & Stuart 1985). Originally described in 
a monotypic genus (Tristram 1880), it has even been considered a 
‘bakkamoena offshoot’ (Peters 1940: 97). 

Otus alius differs from O. [magicus]| sulaensis (see Plate 3, lower right; 
Sula Islands, n=2) in its much more extensively feathered tarsus; less 


P. C. Rasmussen 148 Bull. B.O.C. 1998 118(3) 


prominent dark eye patch; less heavily barred auriculars and spotted 
lower throat; shorter bill (Table 1); more uniformly barred brown 
dorsum lacking spots and streaks; more evenly spaced bands on outer 
webs, and light banding on inner webs of outer primaries; scapular 
spots lacking large black central patches; more prominently banded 
tertials and upper tail, and smaller size. Judging from its highly 
distinctive vocalizations and morphology, sulaensis is probably a 
separate species (Hartert 1898, Finsch 1898, Coates & Bishop 1997, 
King 1997) but only two adult specimens exist. 

For three other taxa from islands off Sulawesi (for which the 
combined world total of adult specimens is only six!), vocalisations are 
unknown and the taxonomy unsettled. From O. [m.] Ralidupae (see 
Plate 3, lower left, TTukangbesi; n=2), alius differs in its darker brown 
colouration; less extensive tarsal feathering; less vermiculated venter; 
less finely patterned eartufts; more prominently banded upper tail; very 
different scapular spots; less strongly banded undersides of primaries, 
but stronger banding of uppersides; and smaller overall size (Table 1). 
From O. [m.] mendeni (Peleng I.; n=3; Eck 1976), alius differs in its less 
vermiculated and unspeckled dorsum (in grey morph); much longer tail 
and especially wing; less streaked and more barred venter; and much 
more extensive tarsal feathering (Table 1). From the unique specimen 
of O. [m.] staoensis (Siau I. in Sangihe Is.; Schlegel 1873), alius differs 
in its much larger size, especially wings and tail; its lack of a 
conspicuous pale ochre collar; its much broader tail banding (‘Table 1); 
and its less richly coloured, more regularly barred venter. 


Otus magicus sensu latu 

Additionally to the characters listed above and below, Otus alius 
differs from most of the remaining races of magicus in having the 
tertials less broadly and prominently banded. It is about the size of the 
two small Lesser Sundas taxa (albiventris and tempestatis—grey morph 
of the latter is shown top right, Plate 3) and smaller than the remaining 
(Moluccan) taxa, samples of which were combined herein, but its feet 
and claws are relatively large for its size (Table 1). From albiventris 
(Lombok to Lomblen; n=25), alius differs in its much shorter, more 
rounded, unstreaked eartufts; shorter, less profuse rictal bristles; and 
breast more uniform with rest of venter. Contra Sibley & Monroe 1993, 
O. alfredi (Flores; n=3) is not the red morph of albiventris, but belongs 
to the O. spilocephalus group (Hartert 1925, Stresemann 1925, Widodo 
et al. unpublished). From grey-morph tempestatis (see Plate, upper 
right, Wetar I.; n=6), alius has venter less mealy; less dark above the 
eye; and more coarsely barred eartufts; and from the red morph of 
tempestatis (n=6), it has narrower pale cinnamon (less orange) barring 
more prominently black-edged below. 

From O. m. magicus (Amboina and Scram; n= 24) alius differs greatly 
in having the dorsum much less heavily spotted pale, and the venter 
with greatly reduced streaks and much less speckled, lacking bold black 
markings; scapulars lack irregular black marks. From Otus m. bouruensts 
(Buru; n=9), alius is darker above, lacks whitish hindcollar, has more 
regular barring on primaries; less narrowly barred tail; paler eyepatch; 


P. C. Rasmussen 149 Bull. B.O.C. 1998 118(3) 


eartufts more barred; scapular spots smaller; and underparts browner, 
lacking strong, sharp streaks. From O. m. leucospilos (Halmahera and 
Bacan, n=23), alius has a less pronounced facial disk; less white 
spotting and more even patterning below, without strong contrast 
between breast and belly; less uniform dorsum, lacking streaks; shorter 
eartufts; and rather different scapular spots; O. m. obira (Obi, n=1 
specimen examined and photos of type seen) resembles a rather dark 
leucospilos. From QO. m. morotensis (Ternate and Morotai, off 
Halmahera; m= 21), alius differs essentially as from leucospilos but is also 
paler. Another form attributed to magicus (Wolters 1970; not 
mentioned by Marshall 1978), O. m. obsti Eck 1973, is probably a 
mislabelled manadensis (Lambert & Rasmussen unpublished). 


Eastern Indian Ocean endemics 

Otus alius differs greatly from O. enganensis (Enggano Island; n=4 
specimens examined, and photos of 3 others) in being less rufous and 
more barred, with less streaking below and less vermiculation above; 
scapular spots smaller with larger, blacker tips; eartufts less finely 
barred; tertials and tail more banded; bill smaller; toes heavier; and 
tarsi less thickly feathered. The rather distinctive calls of enganensis 
suggest it is a separate species (van Marle & Voous 1988, King 1997), 
and this is here considered warranted as its plumage is very unlike 
magicus. Otus alius differs from the much smaller (but geographically 
nearest) O. umbra (Simeulue I.; n=2) in its less rufescent plumage 
overall; paler facial disk; barred eartufts; larger black tips to whiter 
scapular spots; broader pale bands on upper tail surface; and 
proportionately much larger feet and claws. The vocalisations of wmbra 
are very distinct (Marshall 1978) so that, despite its similarity in 
plumage (but not size and proportions) to enganensis, its specific status 
is not in doubt. A closer relationship of alius with enganensis and umbra 
than with magicus is suggested by their shared wing formulae (Table 1; 
unpubl. data) and bill colouration, qualitatively unlike those of 
all Wallacean magicus subspecies, as well as sulaensis, beccari, and 
insularis. Although O. mentawi (Mentawai Is.; n=7 specimens 
examined, photos of another) has been thought a member of 
the magicus group (Neumann 1939, Eck & Busse 1977), it is much 
closer to O. bakkamoena (Riley 1929, Deignan 1950) and is treated as 
a separate species by most on the basis of its highly distinctive 
vocalizations. 

Marshall (pers. comm. 1997) thought the Nicobar Scops-Owl must 
be the same taxon as a pair of owls that apparently bred in 1976 on 
Pulo Perak, off N.E. Sumatra, and the latter were listed in Otus 
magicus in Marshall & King (1988), and as ‘Small-Island Scops Owl 
Otus sp.’ (Wells 1983). However, photos of two of the Perak owls 
taken by M. Avery (no specimens exist) show them to be very 
different from all these taxa, differing from alius in their long, 
pointed, speckled eartufts; heavily streaked forehead and underparts; 
much white on lower underparts; both webs of scapulars largely 
white with small triangular black tips; white feathering on thin legs, 
with distal third bare; and relatively much smaller feet and claws. 


P. C. Rasmussen 150 Bull. B.O.C. 1998 118(3) 


The Pulo Perak birds show no salient differences in plumage from 
the type of Otus sunia nicobaricus. 


Other taxa 

From the variable O. manadensis (Sulawesi; n=47), alius differs by 
the same plumage and proportional characters as those by which it 
differs from the Lesser Sundas magicus forms, except that the banding 
of the tertials is similar; alius has the breast more similar to the lower 
underparts; lacks black spotting and pale rufous bases to eartufts; has 
auriculars paler; usually broader pale tail bands; paler central 
forecrown; larger claws; and smaller rounded white scapular spots. 
Sangihe Island Otus (n=3) have been thought identical to manadensis 
(Sharpe 1875, Meyer 1884, Finsch 1898), and differ essentially from 
alius as does manadensis, they will be dealt with elsewhere (Lambert & 
Rasmussen unpublished). 

From Japanese and Taiwanese taxa (O. e. elegans, Ryukyu and 
Amami Is.; n=15; botelensis, Lan Hsu I|.; n=1 specimen, photos of 6 
additional specimens and one living bird examined; and interpositus, 
Borodino Is.; no specimens examined but photos of 6 specimens 
seen) alius differs greatly in its wing formula; shorter wings; less finely 
vermiculated but more uniformly coloured plumage: shorter, barred 
eartufts lacking rufous inner webs; broader pale tail bands; and lack of 
small black markings in scapular spots. From O. [elegans] calayensis 
(Batanes and Calayan, Philippines; n=4), a few individuals of which 
alius somewhat resembles due to their barred mantles, it differs in 
being browner; with barred eartufts lacking rufous edges; less heavy, 
less finely barred tarsal feathering; less conspicuous streaking and 
more tricoloured barring below; less uniform uppertail surface and 
tertials; tail shorter; and less uniform, less ochraceous facial disk. 
From the Philippine taxa of O. mantananensis (cuyensis, n=3; 
romblonis, 5; sibutuensis, 13; steerer, 1; and mantananensis, 2 specimens 
examined and photos of another), aliuws has narrower breast streak- 
ing; more tricoloured barring of venter; unstreaked lower throat; 
and prominently banded upper tail and tertials (similar to the 
nominate form); there are additional differences from each individual 
taxon as well. None of the African taxa in the manadensis superspecies 
are as similar to alius as are some of the taxa treated above, and hence 
they are not dealt with here, although all were examined for this 
study. 


Life history data 

The holotype had greatly enlarged testes (26 X 14mm) when 
collected on 3 March. The paratype was noted to have the ‘ovary 
granular’ but an accompanying drawing showed it to be approaching 
breeding condition (ca. 10mm long, largest follicle ca. 1 mm) on 2 
April, although it was moulting the inner primaries (P8—P10) at the 
time. The holotype, collected as it flew over a clearing, had eaten ‘a 
spider and a beetle (Apogonia ferruginea F.) (Abdulali 1967: 172). The 
paratype ‘was found perching singly on a tree, ca. 50 ft above the 
ground in the coastal forest, ca. 1 km away from the water line ...’ 


P.C. Rasmussen isi Bull. B.O.C. 1998 118(3) 


(B. Biswas in litt. to J. T. Marshall, 24 April 1979); its stomach 
‘contained a mangled 4 inch gecko’ (Abdulali 1978: 759). 


Voice 

The call of the female was rendered by S. S. Saha as ‘ooo-m’ 
(Abdulali 1978: 759) or as ‘rising long drawn (lasting for 2 to 2.5 s), 
single syllable melancholic moan, repeated after 3 to 5s, ‘otin’ 
pouring out its steady moaning note continuously for more than 30 
minutes or so... at about 2000 hrs ...’ (B. Biswas im litt. to J. T. 
Marshall, 24 April 1979). Territorial calls of the male (those used as 
taxonomic characters by Marshall 1978) are unknown. 


Conservation 

Since only two specimens are known, from the same place on one 
island, it may be that alius is at some degree of risk. Its possible 
restriction to Great Nicobar is suggested by the fact that it was not 
encountered on the more northerly Nicobars despite the more extensive 
collecting done there (Hume 1873, Butler 1899-1900, Richmond 1902). 

Prior to collection of the first alius, the only collecting on Great 
Nicobar was apparently 2—3 days by A. O. Hume’s party (Hume 1873) 
and under a month by W. L. Abbott (Richmond 1902). However, the 
Great Nicobar group includes the nearby Little Nicobar (Stattersfield 
et al. 1998), which is also little-known, so alius may occur there, and 
surveys are urgently needed to establish the status of alius. In any case, 
now that this well-marked, highly isolated taxon has finally received a 
name, the process of studying and conserving it can commence. 


Acknowledgements 


Special thanks go to J., T. Marshall, Jr., who shared his slide collection and notes, and 
gave his blessing for the preparation of this description; to B. F. King, with whom 
discussions helped form the early impetus for this study; and to H. Abdulali. Others who 
provided access to collections, loans, assistance, discussion, and/or read the manuscript 
include N. J. Collar and F. R. Lambert, BirdLife International; M. LeCroy, P. Sweet 
and G. F. Barrowclough, American Museum of Natural History (AMNH); L. Joseph, L. 
Bevier and D. Agro, Academy of Natural Sciences of Philadelphia (ANSP); M. P. 
Walters and R. P. Prys-Jones, BMNH; A. Rahmani and S. Unnithan, BNHS; G. Hess, 
Delaware Museum of Natural History (DMNH); D. Willard, Field Museum of Natural 
History (FMNH); R. A. Paynter, Jr., Museum of Comparative Zoology, Harvard 
University (MCZ); C. and J.-F. Voisin, Museum National d'Histoire Naturelle, Paris 
(MNHN); S. Eck, Staatliches Museum ftir Tierkunde, Dresden (MTD); R. Dekker, 
Nationaal Natuurhistorisch Museum, Leiden (RMNH); L. Vijayan and R. Sankaran, 
Salim Ali Centre for Ornithology (SACON); R. B. Payne and J. Hinshaw, University of 
Michigan Museum of Zoology (UMMZ); R. S. Symonds, University Museum of 
Zoology, Cambridge (UMZC); G. R. Graves, S. L. Olson, E. A. Schreiber and R. C. 
Banks, National Museum of Natural History, Smithsonian Institution (USNM); D. 
Allen and the authorities of the Yamashina Institute, Tokyo (YJ; collection not visited); 
S. Frahnert and J. Fiebig, Museum fiir Naturkunde, Berlin (ZMB); K. Roselaar, 
Zoologisch Museum, University of Amsterdam (ZMA); S. S. Saha, Zoological Survey of 
India (ZSI); M. D. Gottfried, Michigan State University Museum; and P. Alstrém. 
Finally, I am especially indebted to L. B. McQueen for painting the Plate. 


References: 
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JF. Bombay Nat. Hist. Soc. 64: 139-190. 


P. C. Rasmussen 152 Bull. B.O.C. 1998 118(3) 


Abdulali, H. 1972. A catalogue of the birds in the collection of the Bombay Natural 
History Society—11. Strigidae and Caprimulgidae. 7. Bombay Nat. Hist. Soc. 69: 
102-129. 

Abdulali, H. 1978. The birds of Great and Car Nicobars with some notes on wildlife 
conservation in the islands. 7. Bombay Nat. Hist. Soc. 75: 744-772. 

Ali, S. & Ripley, S. D. 1969. Handbook of the birds of India and Pakistan. Vol. 3. Oxford 
Univ. Press, Bombay. 

Ah, 5S. & Ripley, oe Ds 1981. Handbook of the birds of India and Pakistan. Vol. 3. Second 
Edn. Oxford Univ. Press, Bombay. 

Butler, A. L. 1899-1900. The birds of the Andaman and Nicobar islands. 7. Bombay Nat. 
Hist. Soc. 12: 386-403, 555-571, 684-696; 13: 144-154. 

Coates, B. J. & Bishop, K. D. 1997. A guide to the birds of Wallacea. Dove, Alderley, 
Queensland. 

Collar, N. J. & Stuart, S. N. 1985. Threatened birds of Africa and related islands. 1.C.B.P., 
Cambridge, U.K. 

Deignan, H. G. 1950. The races of the Collared Scops Owl, Otus bakkamoena Pennant. 
Auk 67: 189-201. 

Eck, S. 1976. Die Vogel der Banggai-Inseln, insbesondere Pelengs (Aves). Zool. Abh. 
Staatl. Mus. Tierk. Dresden 34: 53-97. 

Eck, S. & Busse, H. 1977. Eulen Die rezenten und fossilen Formen. Aves, Strigidae. Neue 
Brehm Bucherei No. 469. A. Ziemsen Verlag, Wittenberg, Germany. 

Finsch, O. 1898. Uber Scops magicus (S. Mill.) und die verwandten Arten. Notes Leyden 
Mus. 20: 163-184. 

Gurney, J. H. 1884. On some eastern owls. Ibis (Ser. 5) 2: 169-173. 

Hartert, E. 1898. List of a collection of birds made in the Sula Islands by William 
Doherty. Nov. Zool. 5: 125-156. 

Hartert, E. 1925. Types of birds in the Tring Museum. B. Types in the general 
collection. VI. Nov. Zool. 32: 259-276. 

Hume, A. O. 1873. The islands of the Bay of Bengal. Stray Feathers 2: 29-324. 

King, B. F. 1997. Checklist of the birds of Eurasia. Ibis, Vista, California. 

Lambert, F. R. & Rasmussen, P. C. ms. Scops-owls of the Sangihe Islands. 

Marshall, J. T. 1972. Review of ‘Handbook of the birds of India and Pakistan together with 
those of Nepal, Sikkim, Bhutan and Ceylon’. Auk 89: 207-209. 

Marshall, J. T. 1978. Systematics of smaller Asian night birds based on voice. Orn. 
Monogr. No. 25: 1-58. 

Marshall, J. T. & King, B. F. 1988. Genus Otus. Pp. 331-336 in: Amadon, D. & Bull, J. 
Hawks and owls of the world: a distributional and taxonomic list. Proc. Western 
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Philadelphia 41: 1-37. 

Meyer, A. B. 1884. Ueber neue und ungentigend bekannte Vogel, Nester und Eier aus 
dem Ostindischen Archipel im K6nigl. Zoologischen Museum zu Dresden. Abh. 
Naturwiss. Ges. Isis, Dresden 1: 1-64. 

Neumann, O. 1939. Six new races from Peling. Bull. Brit. Orn. Cl. 59: 104-108. 

Peters, J. L. 1940. Check-list of birds of the world. Vol. 4. Harvard Univ. Press, 
Cambridge, Mass. 

Richmond, C. W. 1902. Birds collected by Dr. W. L. Abbott and Mr. C. B. Kloss in the 
Anduman and Nicobar islands. Proc. U.S. Nat. Mus. 25: 287-314. 

Riley, J. H. 1929,. A review of the birds of the islands of Siberut and Sipora Mentawi 
group (Spolia Mentawiensia). Proc. U.S. Nat. Mus. 75: 1-45. 

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Press, Bombay. 

Schlegel, H. 1873. Revue de la collection des oiseaux de prote faisant partie du Musée des 
Pays-Bas. Aves Noctuae. Oti. Tome II, Monogr. 11. E. J. Brill, Leiden. 

Sharpe, R. B. 1875. Catalogue of the birds in the British Museum. Vol. Il. Striges. 
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Watson, J. 1980. The case of the vanishing owl. Wildlife 22: 38-39. 

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Address: Pamela C. Rasmussen, Room 336 NHB MRC 114, Smithsonian Institution, 
Washington, D.C. 20560-0131, U.S.A. 


© British Ornithologists’ Club 1998 


A case of ‘song-capture’: Rufous-naped Lark 
uses nightjar song in place of its own song 


by Jennifer F. M. Horne & Lester L. Short 
Received 8 March 1997 


Cases of mimicry involving song are common in birds. These usually 
involve the learning and incorporation into its own song by one species 
of part or all of the song of another species; sometimes the mimicked 
song is taken as one of several songs in a species’ repertory, as in some 
emberizine finches (Baptista et al. 1981, Petrinovich & Baptista 1987, 
Baptista 1988). Often these finches improvise and can acquire and 
employ parts of the songs of other finches, as of part of a Strawberry 
Finch Amandava amandava song and modified Lincoln’s Sparrow 
Melospiza lincolni songs by White-crowned Sparrows Zonotrichia 
leucophrys (Baptista & Morton 1988). Less frequent is the acquisition 
by one species of the full song of another that is then used exclusively as 
if it were the singer’s own song. A captive Gouldian Finch Chloebia 
gouldiae learned and mimicked all but a terminal element of the song of 
an adjacently caged Strawberry Finch (Baptista 1973) as its sole song. 
Short (1966) documented the singing by a wild Field Sparrow Spzzella 
pusilla of the typical trill song of a Chipping Sparrow S. passerina as the 
sole song of the Field Sparrow. All known examples involve related 
con-familial species, or species of the same order. We here document 
a case of such ‘song-capture’ involving the widespread African 
Rufous-naped Lark Muirafra africana. This lark typically has a 
relatively simple song uttered from the ground, or a perch (on a rock, 
fence or tree), and an aerial song that is more complex; the latter, but 
not the former, often contains mimicked bits from vocalisations of 
other birds. In some areas, such as our honeyguide study site in central 


J. F.M. Horne & L. L. Short JS): Bull. BOC: A998 18GB) 


Kenya, the aerial song is rarely employed by the Rufous-naped Lark, 
perhaps due to sympatry with its more common congeners the Flappet 
Lark M. rufocinnamomea and especially the abundant Fawn-coloured 
Lark M. africanoides, both favouring repetitive aerial songs heard for 
many months of the year. The Rufous-naped Lark song involved here 
is its ground or perch song; and the song captured, that is, learned and 
sung consistently, exclusively and almost perfectly copied, is that of the 
common Montane Nightjar Caprimulgus poliocephalus. 

The site is a degraded hilly woodland at 1,850 m just north of the 
equator and 25 km east of Lake Baringo (thus in highlands just east of 
the Rift Valley) on Ol] ari Nyiro Ranch. At the site, that of our 
permanent camp, Montane Nightjars had sung nightly during late 
April 1994, and in parts of all preceding years back to 1983. As in all 
nightjars of the genus Caprimulgus known to us, the Montane Nightjar 
sings only in hours of darkness, and the short periods of near darkness 
about dusk and dawn. ‘The Rufous-naped Lark occurs only sporadically 
about the site, which could be a factor in acquisition of an allospecific 
song. Males sing from the ground or a perch at intervals throughout 
the day, but not so early or so late as we hear aerial songs of the 
Fawn-coloured and Flappet larks. 

At 13.51 hrs on 30 April 1994 we heard 7 or 8 Montane Nightjar 
songs from the slope above camp. The sun shone at the time, although 
the sky was partly overcast with some nimbus clouds in view. From 
14.07 to 14.14 hrs we again heard these songs at c. 3 per minute. This 
continued off and on to 14.30 hrs, when they came from close at hand, 
ceasing at the onset of rain at 14.43 hrs. Other chores kept us from 
seeking the diurnally singing “‘nightjar’ that day, although we 
recognised the unusual occurrence of diurnal singing (in East Africa we 
have not heard Montane Nightjars sing after 06.15 hrs or before 
18.25 hrs). Prior to the rain we had sent an assistant to seek the singer, 
but he reported that the songs ceased when the bird was approached. 
At 16.13 hrs, following the rain, the nightjar songs commenced, 
continuing to 16.47 hrs at some distance, and from 16.48 to 16.52 hrs 
from closer to us. Joining the search we flushed a Yellow-necked 
Spurfowl Francolinus leucoscepus that landed at a point near the singing 
nightjar, but no nightjar or other bird flew up. Further songs came 
from upslope to the northwest at 16.54, 17.01 and 17.08 hrs, then 
sporadically to 17.50 hrs. There were three more of the songs between 
18.00 and 18.13 hrs, and another three to 18.30 hrs. ‘They ceased at that 
time. By 19.05 hrs it was totally dark. Then from 19.30 to 20.00 hrs the 
apparently identical song emanated from farther upslope. 

Diurnal ‘Montane Nightjar songs’ (Fig. 1; rendered variously by us 
as peea-leeur or feea-feeurr, and by Fry et al. 1988 as pee-yay-yoh, 
pee-yarrh) thereafter were heard sporadically when we were at camp 
during the day (mainly afternoons when, after visiting one or another 
of our 16 honeyguide study sites, we were back for chores and 
note-writing). On 1 May JH was able to seek the singer, going upslope 
behind camp at 11.00 hrs; she tape-recorded several of the songs, 
played back what she had recorded, and verified that a Rufous-naped 
Lark was the source of the ‘nightjar’ songs. The rest of us (LS, assistant 


J. F. M. Horne & L. L. Short 155 Bull. B.O.C. 1998 118(3) 


yo -F HD @ 


8 
6 ‘ 

4 wom TS, 
2 


FREQUENCY IN KILOHERTZ 
hw. Se “o>. 160 


TIME IN SECONDS 


Figure 1. Sonograms of: (A) song of Rufous-naped Lark; (B) nightjar-like song of a 
Rufous-naped Lark; (C) song of Montane Nightjar (all from central Kenya, last one 
courtesy of Library of Natural Sounds, Laboratory of Ornithology, Ithaca, New York). 


Stephen Njagi and technician the late Dickson Chepus) confirmed that 
the lark responded to the playback of ‘its’ song with more ‘nightjar’ 
songs. We continued to hear the diurnal ‘nightjar’ songs thereafter on: 
2 May after 14.20 hrs, 3 May in mid-afternoon, 4 May in the morning, 
5 May in the afternoon, 7 May in the afternoon (by this time, when and 
as we could, we put nets up about small bushes from which it had 
sung), and 8 May in the afternoon, when assistant Njagi observed it for 
a half hour, and assured us that all of the nightjar songs emanated from 
the lark. We then verified that the singer indeed was a Rufous-naped 
Lark, which by now we referred to as the ‘nightjar lark’. We heard no 
further diurnal nightjar songs during May, and were busy with the 
honeyguides in their main breeding period. LS then returned to. New 
York, and JH continued the studies through June. On 24 June the 


J. F.M. Horne & L. L. Short 156 Bull. .B.O-€, 19981 Va3p 


assistant reported a ‘nightjar lark’ singing near another honeyguide site 
3 km northeast of the site at camp, on a degraded, formerly wooded 
slope (now grassy after April and May rains) with scattered bushes and 
small trees. JH verified that the singer of the nightjar-like song was 
a Rufous-naped Lark, presumably the same male (or, conceivably, a 
brother, or, assuming reproduction of such a singer, a son, or its 
father), and that it sang only the ‘nightjar’ song. On subsequent days at 
this site the Rufous-naped Lark continued incessantly, singing only 
‘nightjar’ songs (25, 26, 28 June). When LS returned in early July plans 
were made to record again the voice of the lark, and to collect it. 

Meanwhile, nocturnal Montane Nightjar songs had ceased in May, 
whereas a few Rufous-naped Larks still sang at several localities, all 
using the typical ‘sweet Mary’ song (sonogram in Fig. 1; rendered 
variously by Keith et al. 1992 as tseep-tseeoo, tiree-tiroo and 
teeoo-teewee, by us as feet-be, pee-oh) at about double the pitch of the 
Montane Nightjar’s song, slightly shorter than that song and with a 
somewhat greater frequency range, but nonetheless a simple, whistled 
song remarkably like that of the Montane Nightjar both structurally 
and to our ears. Montane Nightjars have no other commonly heard, 
loud vocalisations, but Rufous-naped Larks also employ: the complex 
aerial song mentioned above, heard only rarely in our area (and heard 
not at all from the ‘nightjar lark’); a single or double-noted, 
ventriloquial whistle uttered from a perch, that we have not heard from 
the lark in question; and a so-called ‘alarm’ call pee-wit or tweekiree that 
we frequently hear (pee-wit version; Keith et al. 1992), and heard from 
the ‘nightjar lark’. 

On 6 July we found the lark still singing ‘nightjar’ songs (and 
exclusively nightjar songs repeatedly) at that second site, singing from 
bushtop perches in a 1-ha area on a slope. We recorded its voice as it 
consistently sang the nightjar-like song at 2 to 5 per min, and we also 
recorded several of its pee-wit calls as we followed it about for over an 
hour. We collected it as it sang the ‘nightjar’ song from a dead bushtop 
located in a ring of bushes barely marking an old ‘boma’ (a temporary 
night camp and protective stock pen, probably from an earlier period of 
the Ranch, but possibly dating back 110 years to the occupation of the 
region by the now extinct Laikipiak clan of the Maasai). ‘The habitat is 
degraded bushed grassland with a 20% coverage of a few trees, mostly 
long-dead Olea europaea, but mainly of bushes, especially Carissa 
edulis, Euclea divinorum and Rhus spp. The lark proved to be a male M. 
africana atht with testes to 8mm; it weighed 47.5 g and had insect 
remains including fragments of ants in the ventriculus. The specimen is 
in the collection of the National Museums of Kenya. 

The singing rates for Rufous-naped Larks given by Keith et al. 
(1992) are misleadingly rapid. We found that the rate of delivery varies 
greatly from 1 or 2 to up to 10 times a minute. Nor are the two parts of 
this lark’s song always included; the first half, swee-ee, is uttered 
sporadically among series of full songs. The lark that sang the 
nightjar-like song occasionally rendered only this first part, and indeed 
Montane Nightjars often interject such a partial song among their 
songs. 


3. F.M. Horne & L. L. Short 157 Bull. B.O.C. 1998 118(3) 


The only additional datum from the field is that our assistant heard 
‘Montane Nightjar’ songs during one day in mid-August 1994, while 
we were away, at a site 2 km from our camp and 2.5 km from the site of 
the ‘nightjar lark’ in July, and over a small hill from both. On his own 
he sought and located the singer, found to be another Rufous-naped 
Lark, singing from a bush in badly degraded bushed grassland about a 
water tank heavily used by elephants, buffaloes, elands, zebra and other 
‘game’, as well as cattle. When we returned in September we could not 
find this lark (possibly a brother of the lark collected; our-assistant is a 
totally reliable observer and keen naturalist with 15 years of experience 
on the Ranch), and in the intervening period up to the present we have 
heard no other diurnal nightjar songs. 

There are no studies of song learning or mimesis in larks of the genus 
Mirafra. Other larks have various songs, some involving complex 
mimicry and incorporation of various sounds (including several 
whistled commands of a shepherd to his dogs) into songs (e.g., Crested 
Lark Galerida cristata; Cramp et al. 1988, 'Tretzel 1965). We note that 
the Rufous-naped Lark’s non-aerial song is relatively simple, without 
mimicked portions, and seems to be readily recognisable as such 
throughout its considerable distribution, even though it may be 
learned. Presumably, during its sensitive period, the ‘nightjar lark’ that 
is the subject of this report completely captured or acquired the full 
Montane Nightjar’s song in or near its natal area, and the song 
underwent no elaboration or modification. It is perfectly possible that a 
male Montane Nightjar had a song post very close to the nest or 
roosting site of the lark when it was young; the nightjar’s habitat 
includes bushed grassland (though it is more in ‘bush’ and more 
wooded sites than usually is the precise habitat of the lark), and a 
Montane Nightjar sings from the same perch, an olive stub, in our 
camp as does the more common Dusky Nightjar Caprimulgus fraenatus, 
but not at the same time. 

Structurally the songs (Fig. 1) are very similar, basically whistles 
with some shifts in pitch. The Montane Nightjar’s song is well below 
the range of frequency of the song of Rufous-naped Larks, but 
congeners of that lark such as the Monotonous Lark M. passerina 
(and others, see sonograms in Maclean 1985) regularly sing lower- 
pitched songs, suggesting that various species in the genus have this 
capability. The exactness of the match is remarkable, as attested by 
our inability to distinguish the lark’s song from the ‘nightjar’s. athe 
lark matched all the rises and falls of the song at the appropriate 
times, and correctly shifted to the trilled ending at the time when a 
nightjar would do so (the trill itself is shorter than the comparable 
trill portion of the nightjar’s, and indeed both parts of the song are 
slightly shorter than in the nightjar, but this does not detract from 
the closeness of the match). 

One certainty is that the nightjar’s song was learned at night. Thus, 
within the lark’s sensitive period the template for learning is open night 
and day. Experimenters thus have the night hours as well as daytime 
for use of playback in learning experiments with at least some larks, and 
doubtless other birds. 


J. F. M. Horne & L. L. Short 158 Bull. B.O.C. 1998 118(3) 


A moot point is whether or not the nightjar’s song employed by the 
lark could have attracted a potential mate for the lark. Probably this is 
unlikely, given the differences between the songs of the two species. 
Perhaps the movement of the lark from our camp site to the other site 
between May and June represented a shift in locale due to the failure of 
the lark to attract a female lark? Certainly no other Rufous-naped Lark 
of either sex was heard or seen at either site. Of course the singers 
possibly were different individuals, as it seems quite likely that two (or 
even 3) males could have been brothers exposed to the same nightjar 
tutor at the same time in their lives. 

‘The paucity of reports of song capture (as opposed to acquisition of 
parts of a song for incorporation into the imitator’s song, or the taking 
of another, even wholly unrelated species’ song as one element in the 
mimicker’s repertory) suggests that it is an unusual if not remarkable 
event in a few avian species. 


Acknowledgements 


We are grateful for support from the National Museums of Kenya and the American 
Museum of Natural History, and to the Gallmann Memorial Foundation for permission 
to conduct research on OI] ari Nyiro Ranch. The study owes much to our field staff, 
especially to fearless, competent Stephen Njagi who daily carries out tasks on long, often 
lonely hikes in the presence of large mammals and other potential hazards that would 
deter most humans. Luis Baptista as ever graciously provided useful information and 
comments on the manuscript. 


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Baptista, L. F. & Morton, M. L. 1988. Song learning in montane White-crowned 
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Baptista, L. F., Morton, M. L. & Pereyra, M. E. 1981. Interspecific song mimesis by a 
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Maclean, G. L. 1985. Roberts’ Birds of Southern Africa, 5th edn. J. Voelcker Bird Book 
Fund. 


Petrinovich, L. & Baptista, L. F. 1987. Song development in the White-crowned 
Sparrow: Modification of learned song. Animal Behav. 35: 961-974. 

Short, L. L. 1966. Field Sparrow sings Chipping Sparrow song. Auk 83: 665. 

Tretzel, E. 1965. Imitation und Variation von Schaferpfiffen durch Haubenlerchen 
(Galerida c. cristata [L.]). Zeitsch. Tierpsych. 22: 784-809. 


Address: Jennifer F. M. Horne & Lester L. Short, Ornithology Dept., National Mus. 
Kenya, P.O. Box 40658, Nairobi, Kenya. 


© British Ornithologists’ Club 1998 


A. #. Begazo & T. H. Valqui 159 Bull. B.O.C. 1998 118(3) 


Birds of Pacaya-Samiria National Reserve 
with a new population (Myrmotherula 
longicauda) and new record for Peru 
(Hylophilus semicinereus) 


by Alfredo 7. Begazo © Thomas H. Valqui 
Received 6 May 1997 


The Peruvian Amazon encompasses a variety of forest ecosystems. A 
relatively small but significant part of these ecosystems is the flooded 
forest. This unique formation has a variety of endemic plants and 
animals (Ayres 1993). In addition, the flooded forest represents an 
important component for the life cycle of fish and a source of food for 
local inhabitants (Padoch 1988). However, this ecosystem is among the 
most vulnerable in Amazonia because of its accessibility for resource 
extraction by water transport. 

The Pacaya-Samiria National Reserve (PSNR) covers approximately 
2,080,000 ha, being the largest protected area in the Amazon basin. The 
overall topography is dominated by a flat terrain subject to inundation 
to varying extents, except for a small area in the southwestern part of 
the reserve. Its strategic location between two major rivers and the 
predominantly flat terrain result in a peculiar mosaic of forest 
formations. As much as 51% of the total area is composed of flooded 
forest, 34% constitutes seasonally flooded forest, and 1% lakes and 
rivers; 13% of the forest is high enough to escape inundation (Malleux 
1975). Seasonally flooded forest and high forest show a similar floristic 
composition (Encarnacion 1985), whereas forest with permanent or 
near-permanent water shows remarkable differences in floristic 
composition and structure. 

As the reserve’s strategic location results in a great variety of forest 
formations, its position also entails being surrounded by a particularly 
high density of people. The second largest human concentration in the 
state of Loreto (after the city of Iquitos) is found along large rivers. 
Such rivers, for the most part, run across prime agricultural land, the 
result of seasonal sedimentation along their banks that make them 
particularly attractive to human settlement. In addition, large rivers 
represent the only means of transport for people and local products. 
Consequently, PSNR is subject to pressure from extractive activities of 
local inhabitants settled on the periphery. These extractive activities 
selectively affect animal species of large size which, despite being under 
hunting pressure, maintain relatively healthy populations within the 
reserve as opposed to extremely low densities or near local extinction in 
the surrounding unprotected areas (Begazo & Bodmer in press). Large 
cracids, psittacids, herons, ducks and tinamous are the groups most 
heavily hunted within the reserve. 

The present manuscript is the result of 6 months of field work 
between 1992 and 1996. Extensive mist-netting and tape-recording 


A. }. Begazo & T. H. Valqui 160 Bull. B.O.C. 1998 118(3) 


73°33} 
4°24" 


ea ea Rio Maranon 


oe 7" 
6°08' 


Figure 1. ‘The Pacaya-Samiria National Reserve. 


were carried out by the authors throughout the reserve at a total of 
35 surveying stations, of which 14 were situated along the Yanayacu 
river, 6 on the Pacaya river, 13 along the Samiuria river, 5 along a 
transect between the Maranon river and Yanayacu river, and 6 along 
the Maranon river including river islands (Fig. 1). At each surveying 
station, mist-nets were placed along freshly cut trails, and observations 
and tape-recordings were made in the vicinity of the mist-net lines 
and along previously existing trails. Transects were located randomly 
in a vertical fashion along the rivers within the reserve and had a 
varying length. Transects had a double purpose; in some cases they 
were used for general recording of vertebrates and plants, and in others 
for censusing game animals. 'Transects of the first kind measured 1 or 


A. }. Begazo & T. H. Valqui 161 Bull. B.O.C. 1998 118(3) 


2km from the river towards the interior of the forest; censusing 
transects measured 4 to 5 km. Altogether, an estimated 400 km were 
covered in surveys within the forest and along rivers throughout the 
reserve. 

We follow the sequence and taxonomy suggested by Parker et al. 
(1996). In addition to our own records, we have drawn on data from 
previous ornithological surveys conducted within the limits of the 
reserve. We do not include all records from other lists. Species not 
observed by us were included only in cases where we obtained 
confirmatory evidence of their presence in the reserve, e.g. from 
personal communication with the author or authors of such works or 
from local inhabitants. A complete, annotated list of all species 
recorded in the reserve can be obtained from the authors on request. 


Species accounts 
ZIGZAG HERON Zebrilus undulatus 


This elusive heron was recorded on three separate occasions. "THV 
observed an individual under overhanging vegetation at the edge of 
slow-flowing black waters of the Pacaya river. AJB observed the species 
on two occasions in a slow-flowing shallow stream in the interior of the 
forest. The habitat agrees with that described by Davis et al. (1980). 
The birds observed by AJB were at the same place and could have been 
the same individual. On a separate occasion, AJB tape-recorded two 
individuals calling at dawn from a small body of water nearly covered 
by surrounding tangled vegetation at 4°24’'S, 73°06'W, just outside the 
limits of the reserve. The birds called for about half an hour only, 
between 5.00 and 5.30 a.m. Given the abundance of water and the 
swampy nature of the predominant habitats in Pacaya-Samiria, the 
species should be favoured with abundant habitat. With its cryptic 
coloration and shy behaviour it is very easily overlooked. 


SLENDER-BILLED KITE Rostrhamus hamatus 


We frequently saw this species, mostly birds soaring at various 
altitudes. The typical silhouette consists of broad wings with the short 
tail well spread, giving the impression of a continuation of the 
secondary wing feathers. One individual was observed incubating eggs 
for three consecutive days between 6 and 9 July. The nest was placed 
on a horizontal limb of an isolated tree (Pseudobombax munguuba) at 
about 18 m from the ground. 


WATTLED CURASSOW Crax globulosa 


This species has not been reported in Peru in the last 35 years. It was 
known to inhabit riparian vegetation and river islands, which happen to 
be the areas where the fastest human growth has taken place. 
Interviews with local people on the banks of the Maranon suggested 
that the species still exists on the north bank. Two birds were shot in 
August 1996 by a local hunter near the village of Nueva Esperanza 
(4°51'S, 75°06’W). In addition, two birds kept in captivity were 


A. }. Begazo & T. H. Valqui 162 Bull. B.O.C. 1998 118(3) 


photographed and their vocalisations tape-recorded. Thorough field 
work is needed to determine the status of this species in Peru. 


RAZOR-BILLED CURASSOW Crax mitu 


This large game bird has been extirpated from most areas 
surrounding human settlements, but in PSNR there are still healthy 
populations (Begazo 1995). A total of 11 birds were observed in the 
census transects. Razor-billed Curassows seemed to be associated with 
forest with damp soil, especially along stream edges, where they scratch 
the ground in search of food. Analysis of the stomach contents of a bird 
shot by a local hunter showed a variety of vegetable matter including 
seeds and tender shoots, as well as small snails. The generalist diet of 
this terrestrial species (Delacour & Amadon 1973) may help explain its 
preference for moist soil. 


SPIX’S GUAN Penelope jaquacu 


Like the above species, Spix’s Guan is much sought after by hunters. 
Within the reserve it is subject to sporadic hunting pressure; near 
human settlements on the periphery it is virtually extinct. Although 
Amadon & Delacour (1973) suggest that this species makes little use of 
the forest floor, 68% of the birds sighted (n=23) were flushed from the 
ground. Information obtained from local people suggests that the 
species breeds during the months December—March. Stomach contents 
of two birds contained seeds of the Huasai palm Euterpe precatoria. 


COMMON PIPING GUAN Ppile pipile 


Like other large Cracids, this species suffers from human 
persecution. Preliminary studies in three human settlements show that 
this is the species shot in the largest numbers. Although little is known 
about its natural history and movements, we suspect that the seemingly 
high encounter rate with hunters is due to the fact that the species is 
highly frugivorous and must keep moving in search of patchily 
distributing fruiting trees. In contrast, the lower encounter rate with 
territorial species such as Crax mitu may be explained by the fact that 
the latter, once hunted, take a long time to replace empty territories. A 
total of 21 Common Piping Guans were observed along water courses 
and in line transects. 


PALE-WINGED TRUMPETER Psop/ia leucoptera 

Groups of 3 to 11 individuals were registered on 9 occasions during 
the census line transects, in forest subject to seasonal flooding. The 
species is still common in the reserve and, to judge from their tame 
behaviour, they are not being hunted. On 23 February, in a location 
outside the limits of the reserve, a group of 6 adult trumpeters was 
accompanied by two young of about three weeks old and at least one 
juvenile, suggesting that the dominant breeding pairs may make 
consecutive breeding attempts in one season. 


BAND-TAILED NIGHTHAWK WNyctiprogne leucopyga 


On 8 June 1995 THV tape-recorded this species at an oxbow lake 
with abundant grass along the edges (5°09’S, 75°06’W). Later, the 


y 2 ae Begazo & T. H. Valqui 163 Bull. B.O.C. 1998 118(3) 


recording was compared with one of the same species from Brazil 
(Hardy et al. 1986). The species is known from the Guianas, Venezuela, 
the extreme east Colombia, and Brazil from Rio Negro and Madeira 
(Meyer de Schauensee 1970). It has also been reported from the Javari 
River, on the border of Peru and Brazil, about 150 km from our study 
site (Hilty & Brown 1986). 


RUFOUS-NECKED PUFFBIRD Malacoptila rufa 


A total of 5 individuals of this little known species were seen by both 
authors separately. Three birds were seen near Santa Elena check point 
(5°14'S, 74°50'W) and one each at surveying stations on the Samiria and 
Yanayacu rivers (5°13’S, 74°13'W; 5°01’S, 74°13’W). Birds observed at 
Santa Elena were flushed from perches about 50 cm from the ground. 
After flushing, they perched at about 3 m above the ground uttering a 
continuous high-pitched but mellow pzzzu. One of the birds flew back 
to a perch near the ground to resume foraging. It spent long periods 
(2-11 minutes) watching its surroundings before changing perch or 
performing a sally in typical Malacoptila fashion. Once it made a sally 
to take an insect from the ground. 


STRIPE-CHESTED ANTWREN Myrmotherula longicauda 


This report represents a new and disjunct population of this species 
found along the foothills of the Andes between 600-1,200 m (Parker 
et al. 1996). This species is fairly common at 4°56'S, 75°05’'W, in an 
area of short vegetation (5—6 m) in which one or two plant species are 
predominant. Such a plant community is found in permanently flooded 
country with stagnant bodies of black water. It was also seen regularly 
accompanying flocks composed of Myrmotherula brachyura, M. 
minitriestt and occasionally M. surinamensis, which occur in a similar 
habitat along lake or river edges with flowing water. Other members of 
such flocks were Thamnophilus amazonicus, Sakesphorus canadensis and 
Hylophilus semicinereus. A manuscript on the taxonomic status and 
ecology of this population is in preparation by the same authors. 


ASH-BREASTED ANTBIRD Myrmoborus lugubris 


Pairs of this species are fairly common on river islands along the 
Maranon river. The habitat agrees with that given for the species by 
Ridgely & ‘Tudor (1994); our records represent a range extension to the 
locality of Maipuco on the Maranon (4°49’S, 75°07’W). 


BLACK-TAILED ANTBIRD Myrmoborus melanurus 


This very local antbird was found on two occasions within the 
reserve. THV found it on the margin of lake Pastococha (5°10'S, 
75°05'W), and AJB at Quebrada Chiric (4°56’S, 75°05’W), a small 
affuent of the Yanayacu river. As suggested by Ridgely & Tudor 
(1996), the species may have been overlooked. During extensive work 
along Quebrada Chiric AJB did not hear or see it during high water in 
early May, but in the following year during the driest month, late July, 
had several encounters with pairs and individuals of this species. Since 
M. melanurus seems to be associated with damp soil found at water 


A. }. Begazo & T. H. Valqui 164 Bull. B.O.C. 1998 118(3) 


edges, fluctuations in water levels may force birds to the interior of the 
forest where they are less likely to be seen by people who for the most 
part travel by water. As the species was only known to occur east of the 
Ucayali river, these records represent a significant range extension. 


ASH-THROATED GNATEATER Conopophaga peruviana 


The first recorded nest of this species was found on 30 July. While a 
trail was being cut in mature primary forest (5°02’S, 74°59’W), a bird 
was flushed from a nest 84cm above the ground in a sapling with 
spreading branches. Careful inspection showed that the cup-shaped 
nest was placed in an already existing accumulation of dead leaves. The 
inner lining was of fibres, and dead leaves were carefully tucked around 
the exterior of the cup. When found, the nest contained two pearly 
white eggs with brown spots, denser at the wide end. The nest was kept 
under observation for 7 days, the male being found on it in the 
mornings and the female in the late afternoon. While incubating, they 
adopted a peculiar cryptic position with the head lowered, so that the 
cap and scapular feathers looked like a dead leaf with the bill as the 
petiole. 


GRAY ELAENIA Myiopagis caniceps 

A species that is probably often overlooked, as it forages high up in 
the forest canopy (Ridgely & Tudor 1994). We found it to be fairly 
common (heard or seen daily in small numbers). It frequently follows 
mixed-species canopy flocks where it constantly utters a characteristic 
call: tui tui tuiz1z, sometimes followed by high-pitched trill (recordings 
at LNS). Birds perched somewhat horizontally on branches; two were 
observed regularly to flap the left wing before sally-gleaning insects 
from leaves. 


YELLOW TYRANNULET Capsiempzis flaveola 

This distinctive tyrannulet was found in the same place (4°56'S, 
75°05’W) in 1995 and 1996. The habitat is the same as that described 
for the Stripe-chested Antwren. Existing information on its geographi- 
cal distribution shows only scattered records in southern and western 
Amazonia (Ridgely & Tudor 1994). AJB made tape-recordings of the 
typical and sometimes persistent contact vocalisations of foraging pairs 
as well as the trill given by individual birds. On 12-15 July 1996 a pair 
was observed repeatedly visiting what appeared to be an unfinished nest 
placed about 3.5m from the ground in a tall shrub, in a man-made 
opening about 5 m from forest edge. 


JOHANNES’S TODY-TYRANT Hemitriccus 1tohannis 

A single individual was heard and seen within the limits of the 
reserve at 5°00’S, 74°58’W. The bird was foraging at about 7 m in viny 
but light vegetation. The same species was tape-recorded outside the 
limits of the reserve along the T‘ahuayo river, an affuent of the Amazon 
at 4°20’S, 71°52’W. This species seems to be rare in areas outside its 
known geographical range. Jose Alvarez (pers. comm.) reports the 
species for Rio Tigre, near the village of Intuto (specimen collected). 


A. F. Begazo S T. H. Valqui 165 Bull. B.O.C. 1998 118(3) 


These records may suggest a continuous but rare occurrence of the 
species in the area between the main geographical distribution and the 
population of southeastern Colombia (Ridgely & Tudor 1994). 


THREE-STRIPED FLYCATCHER Conopias trivirgata 


Between 21 and 29 June 1995, a group of 5 individuals appeared near 
our house at Nueva Esperanza (4°51'S, 75°06’W), on the south bank of 
the Maranon river. They were identified as C. trivirgata on the basis of 
their small size, and the dark wings in contrast to the colour of their 
backs. The group was vocally so conspicuous that we doubt that the 
birds were there for the previous month that we had been at the site. A 
peculiar behaviour consisted of frequent chases among themselves, 
accompanied by loud and persistent calls both from the birds involved 
in the chase and from other members of the group. After 9 days of 
continuous observation, the group was never seen again. While the 
birds were around, they foraged in old second growth of varzea forest 
and frequently flew across agricultural land to reach isolated trees. This 
observation, and the wide scatter of isolated records of the species in 
Amazonia (Ridgely & Tudor 1994), may suggest some type of 
nomadism or even migration. 


TROPICAL GNATCATCHER Polioptila plumbea 


This widespread but uncommon gnatcatcher was seen following 
canopy mixed-species flocks. The purpose of mentioning this species 
on this account is to highlight the remarkable differences in 
vocalisations between birds in this locality and those of the upper 
Maranon river valley (Hacienda Limon) and west side of the Andes in 
the Department of Piura (pers. obs.). This supports Ridgely & Tudor’s 
(1989) suggestion that two species may be involved. Recordings of 
these vocalisations are deposited at Cornell LNS. 


GRAY-CHESTED GREENLET Hylophilus semicinereus 


The Gray-chested Greenlet’s hitherto known geographical distri- 
bution includes southern Venezuela, French Guiana (Ridgely & Tudor 
1989), and extreme northwest Bolivia (Parker et al. 1996). We found it 
to be common but localised in the Pacaya-Samiria reserve. Birds were 
found in forest edge and in short plant communities growing in 
permanently flooded areas. Our evidence consists of extensive 
tape-recordings and a male collected and deposited in the bird division 
of Peru’s Museo Nacional de Historia Natural Javier Prado. This is the 
first record of the species for Peru. 


Acknowledgements 


The authors want to thank the Biodiversity support Program, The Chicago Zoologica 
Society and the Centro de Datos para la Conservation de la Naturaleza for the financial 
and logistic support provided. We also thank Dr Richard Bodmer for his support during 
part of the field work at the Pacaya-Samiria National Reserve. 


References: 

Ayres, J. M. 1993. As matas de varzea do Mamirarua. MCT-CNPa, Brazilia. 

Begazo, A. & Bodmer, R. in press. Use and conservation of the Cracidae in the Peruvian 
Amazon. Oryx. 


A.L. Archer & D. B. Iles 166 Bull. B.O.C. 1998 118(3) 


Begazo, A. J. 1995. Use of the Cracidae in the Peruvian Amazon. In Proceedings of the 
II International Meeting on Use and Conservation of Amazonian Wildlife, May 
1995. Iquitos, Peru. 

Davis, W. F., Donahue, P. & Perkins, E. G. 1980. Observation of the behavior of the 
Zigzag Heron (Zebrilus undulatus). Condor 82: 460-461. 

Delacour, J. & Amadon, D. 1973. Curassows and Related Birds. American Museum of 
Natural History, New York. 

Encarnacion, F. 1985. Introduccion a la Flora y vegetation de la Amazonia Peruana: 
Estado actual de los estudios, medio natural y ensayo de una clave de determinacion 
de las comunidades vegetales en la Ilanura Amazonica. Candollea (Conservatoire et 
Jardin Botanique de Genéve) 40: 237-252. 

Hardy, W., Coffey, B. & Reynard, G. B. 1986. Voices of the New World Nightbirds, 
Owls, Nightjars, and their Allies. Ara records. Gainesville, FL. 

Hilty, S. L. & Brown, W. L. 1986. A Guide to the Birds of Colombia. Princeton Univ. 
Press. 

Malleux, J. 1975. Mapa forestal del Peru. Memoria explicativa. Lima 161 p. (Mapa escala 
1:1 000 000). 

Meyer de Schauensee, R. 1970. A Guide to the Birds of South America. Livingston 
Publishing Company, Westwood, PA. 

Padoch, C. 1988. People of the floodplain and forest. Pp. 127-141 in J. S. Denslow & C. 
Padoch (eds), People of the Tropical Forest. Univ. California Press. 

Parker, T. A., Stotz, D. & Fitzpatrick, J. W. 1996. Ecological and distributional 
databases for neotropical birds. Jn B. F. Stotz, T. A. Parker III, J. W. Fitzpatrick & 
D. K. Moskovits (eds), Neotropical Birds: Ecology and Conservation. Univ. Chicago 
Press. 

Ridgely, R. & Tudor, G. 1989. The Birds of South America, Vol. 1. The Oscine Passerines. 
Univ. Texas Press. 

Ridgely, R. & Tudor, G. 1994. The Birds of South America, Vol. 2. The Suboscine 
Passerines. Univ. Texas Press. 


Addresses: Alfredo J. Begazo, Department of Wildlife Ecology and Conservation, 
University of Florida, Gainesville, FL 32611, U.S.A. Thomas H. Valqui, 
Department of Ecology and Evolutionary Biology, Princeton University, Princeton, 
NJ 08544-1003, U.S.A. 


© British Ornithologists’ Club 1998 


New bird records for Unguja (Zanzibar 
Island) 


by A. L. Archer & D. B. Iles 
Received & Fune 1997 


In his foreword to R. H. W. Pakenham’s ‘“‘The Birds of Zanzibar and 
Pemba” (1979), Dr J. F. Monk remarks “Tourism, and the limited 
opportunity this offers for scientific observation, is beginning to be 
encouraged and it is possible that in the next decade new data will be 
collected which may revise the status of some species given here’. This 
indeed has proved to be the case; with former restrictions on human 
movement among and within the islands having been _ lifted, 
comparatively easy access to all habitats is now possible. 

Pakenham (1979) lists 183 species known from Zanzibar. Of these, 
109 were known to breed on Zanzibar or outlying islets, the remainder 


A.L. Archer & D. B. Iles 167 Bull. B.O.C. 1998 118(3) 


being migrants or vagrants. The status of many of these species has 
now been revised with the new knowledge obtained over the past five 
and a half years and a further 34 species have been added to the list. Of 
these 14 are palaearctic migrants, 12 African mainland species, seven 
oceanic and one intra-African migrant. 

Besides the two authors, Peter and Ursula Koehler contributed new 
records to the avifauna, their work concentrating mainly on Chumbe 
island, 12 km SSW of Zanzibar town, where they spent time during 
January of 1993 and 1994, ALA has visited Zanzibar on over thirty 
occasions between June 1990 and March 1996, spending 14 months on 
the island. DBI worked in Zanzibar between October 1993 and April 
1995. 

This paper deals with new records for Unguja and the surrounding 
islets. Unguja, the correct name for Zanzibar island, together with 
Pemba island 45 km to the north, makes up the former Zanzibar 
Republic, now part of Tanzania. ‘Zanzibar’, in this paper, refers to 
both Pemba and Unguja. Prior to about 1,900 bird records and 
specimens from the mainland coast nearby were often labelled 
‘Zanzibar’, thus causing some confusion. Stone town 1s part of Zanzibar 
town, situated on the west coast of Unguja. 


Species previously unrecorded 
SHY ALBATROSS Diomedea cauta 


One was killed by fisherman off Matemwe beach, north east Ungyja, 
23 November 1994. The head was brought to ALA by Ali A. Mwinyi, 


Conservation Officer, and is now preserved in the Zanzibar Museum. 


MASKED BOOBY Sula dactylatra 


Although known from Latham island, an uninhabited island 120 km 
south east of Unguja, the first record for Unguja was made by DBI on 
22 July 1995 when an immature was seen being harassed by a Fish 
Eagle Haliaeetus vocifer over the shore of Chumbe island, 12 km 
southwest of Stone town. Two others were seen moving south on the 
same day, and another, also near Chumbe, was observed on 3 February 
1995. 

On 26 November 1995 ALA identified an adult with a broken wing 
tip, caught on Mnemba island, off the north east coast of Unguja. The 
bird was later released after recovery. 


FRIGATEBIRD Fregata sp. 


A single bird observed near Bawe island, Zanzibar bay, by DBI on 
17 December 1994, was immature and could have been either a Greater 
F.. minor or Lesser Frigatebird F. ariel. 


BLACK-HEADED HERON Ardea melanocephala 


ALA observed a single bird in breeding plumage on Bawe island on 
23 January 1993. Subsequently on 8 July 1993 he saw two near 


A.L. Archer & D. B. Iles 168 Bull. B.O.C. 1998 118(3) 


Ngomani and three others near Jongowe, both villages on Tumbatu 
island lying off the north west coast of Unguja. 


OPEN-BILLED STORK Azastomus lamelligerus 

Apart from six seen at Msuka Bay, Pemba, in January 1922, there 
were no other records for the islands until one was seen by ALA on 
10 June 1991, flying near Kibandiko island towards the Unguja coast; it 
was being pursued and mobbed by several Indian House Crows Corvus 
splendens. On 27 January 1992 he recorded another flying 2 km north of 
Bububu, Unguja. DBI saw one flying low along the north coast of 
Chumbe island towards Unguja 2 April 1995 and another was seen at 
Bwawani swamp, Stone town, on 5 November 1995. 


CROWNED CRANE Balearica regulorum 


On 26 June 1992 one with a broken leg was brought to the aviary at 
the Zanzibar Museum. It was one of four which had arrived in a rice 
field east of Masingini forest. Local hunters had been quickly alerted 
and all four were shot, the one survivor being taken to the museum 
aviary where it died a month later. 


HOTTENTOT TEAL Anas hottentota 


This duck has only been recorded from Bwawani swamp. First 
observed by ALA on 23 June 1990 when a pair was seen; then four on 
3 November 1990. Subsequently both authors encountered these ducks 
on nearly every visit to the swamp, with a maximum of 16 on 3 October 
1993. Successful breeding was recorded in July and August with five, 
six and seven ducklings observed. ‘The increase in numbers and the 
presence of well-grown ducklings could be the result of the major 
reduction in the number of Indian House Crows which used to infest 
the swamp area. Despite careful inspection of numerous other ponds on 
the island, no others have been seen. 


EURASIAN MARSH HARRIER Circus aeruginosus 


A male seen soaring near Charawe, east of Jozani forest by ALA and 
D. A. Turner on 4 November 1991 was the first record for Unguja. On 
5 November 1993 DBI saw a female at Bwawani swamp. 


AFRICAN HOBBY Falco cuvieri 


Several sightings of this falcon have been made by both authors, the 
first on 1 November 1990 by ALA at Bwawani swamp. Subsequent 
sightings have been made at Pangeni Ranch, Chumbe island and 
Unguja Ukuu, a village south east of Stone town. A pair was seen in 
flight display by DBI near Unguja Ukuu, indicating possible breeding. 


PURPLE GALLINULE Porphyrio porphyrio 


First observed on Unguja, 3 November 1991 when two were 
observed by ALA, at Bwawani swamp, where they were seen for the 
next four days. Further sightings of one or two were made by both 
authors up to 8 February 1995. An adult, with a partially grown chick, 


A.L. Archer & D. B. Iles 169 Bull. B.O.C. 1998 118(3) 


were observed on 14 July 1992 and again on 25 September 1992 when 
the immature was about full grown. 


STRIPED CRAKE Porzana marginalis 
A single female or possibly immature seen at Bwawani swamp by 


ALA and D. A. Turner on 5 and 7 November 1991. 


LITTLE RINGED PLOVER Charadrius dubius 


A flock of c. 15 was observed at rest on open ground alongside the 
Dongwe/Michamvi road, east of Stone town, by ALA on the evening of 
21 March 1995. 


LESSER (PACIFIC) GOLDEN PLOVER Pluvialis fulva 


Two coming into breeding plumage seen by ALA on 10 April 1993 
on Bawe island. 


MARSH SANDPIPER Tvinga stagnatilis 


A single was seen by ALA at Bwawani swamp on 23 August 1991 and 
again at the same locality on 6 November 1991. Alan Tye (pers. comm.) 
recorded one on tidal flats at Mbweni, 4 km south of Stone town, on 
21 January 1995. 


RUFF Philomachus stagnatilis 


ALA and D. A. Turner observed a single bird at a “fresh water’’ 


drain flowing into the creek near Livingstone House, Stone town on 
5 November 1991. 


ARCTIC SKUA Stercorarius parasiticus 


A subadult light or barred morph was washed ashore at Chumbe 
island following a heavy storm on 14 March 1993. It was released after 
being photographed (Koehler & Koehler 1994). 


POMARINE SKUA Stercorarius pomarinus 


A second winter light morph was captured on the shore of Chumbe 
island on 18 January 1993, again after a heavy storm. It was 
photographed and released only to be killed by a cat (Koehler & 
Koehler 1994). DBI observed an immature chasing terns off Chumbe 
island on 3 January 1994. 


BLACK-HEADED GULL Larus ridibundus 
ALA recorded 11 on 23 January 1992 at the abandoned fresh 


water swimming pool adjacent to Bwawani swamp, Stone town. On 
3 February, the same observer counted 60 on, or flying over, the tidal 
mudflats close by. DBI noted a single immature which frequented 
Stone town harbour between 17 December 1993 and 17 February 1994. 

It is surprising that there are no other records of this gull which has 


expanded its range in eastern Africa dramatically since 1971 (Britton 
et al. 1980). 


Aw. Archer & D) B. Iles 170 Bull. B.O.C. 1998 118(3) 


COMMON NODDY Anous stolidus 

Although known from Latham island in thousands, no records 
existed for Unguja until Peter and Ursula Koehler observed one, two, 
one and four between 9 and 25 January 1993, together with mixed 
flocks of terns off Chumbe island. DBI recorded c. 500 among a 
concentration of c. 3,000 terns seen from the ferry while crossing 
between Dar es Salaam and Zanzibar on 17 December 1994. 


LITTLE/SAUNDERS’S TERN Sterna albifrons/saundersi 


With the lack of specimen material it is possible both species occur. 
Because of their great similarity in non-breeding. plumage and their 
overlap in range, we have combined the two for the purpose of this 
paper. In their present numbers it is inconceivable that R. H. W. 
Pakenham, J. H. Vaughan, D. A. Baird and other observers would have 
missed these distinct small terns. Although Pakenham’s (1979) 
references extend to 1977, little ornithological work was done on the 
islands after the 1964 revolution. It therefore seems safe to assume that 
the terns are a comparatively recent addition to the island’s avifauna. 
The first record by ALA was of c 10 on Bawe island on 6 February 
1991, followed by c. 45 and c. 90 on 23 January and 6 November 1993 
respectively at the same place, c. 100 on Mnemba island on 6 April 
1994, and c. 80 near Jambiani on the east coast of Unguja on 22 March 
1995. DBI recorded large numbers resting on the beach at Paje, east 
Unguja, on 10 December 1994. 


SOOTY TERN Sterna fuscata 

This species breeds in tens of thousands at Latham island but was 
unrecorded in Unguja until 9 and 11 September 1994 when DBI and 
ALA saw two over a Roseate Tern S. dougalli breeding colony off 


Chumbe island. DBI had a further sighting of six among mixed tern 
flocks on 10 November 1994. 


EURASIAN CUCKOO Cuculus canorus 

‘Two cuckoos believed to be of this species were seen by ALA near 
Kizimkazi‘on 12 April 1993. On 7 April 1994 two or three were seen 
near Kiini, north Unguja. One observed at close range showed narrow 
black bars on the underside of the tail and lacked a yellow base to 
mandible, separating it from the African Cuckoo C. gularis. On 9 April 
1995 DBI saw a grey cuckoo in flight near the seashore at Chukwani, 
south of Stone town, which was thought to be this species. 


EURASIAN NIGHTJAR Caprimulgus europaeus 
Although not listed by Pakenham (1979), he noted that Cabanis 
recorded one taken in Zanzibar town in February 1876 and clearly 


identified. DBI netted and ringed a male on Chumbe island on 
28 February 1994. 


EURASIAN SWIFT A pus apus 


At least 2,000 were observed passing Chumbe island on 16 January 
1993 (Koehler & Koehler 1993). On 23 January 1993 ALA and N. E. 


A. L. Archer & D. B. Iles 171 Bull. B.O.C. 1998 118(3) 


Baker saw 40-50 swifts, which were probably this species, overflying 
Bawe island. 


MOTTLED-THROATED SPINETAIL Telecanthura ussheri 

Only recently recorded from Pemba (Archer & Turner 1993), this 
species has now been found to occur on Ungyja. On 13 June 1995 a pair 
was seen by ALA at Fumba and on the same day one was observed at 
Bweleo, both sites on the Fumba peninsula, south of Stone town. DBI 
recorded one over Chumbe island on 3 August 1995 in association with 
Little Swifts Apus affins and two over Jozani forest tree nursery, 
Unguja, on 15 August 1995. ALA saw two birds there on 2 September 
1995. 


CARMINE BEE-EATER Merops nubicus 


The only record for Unguja is of a single bird photographed perched 
in a Casuarina tree outside Jozani forest by Mrs Becky Browning in 
January 1991. 


SAND MARTIN Riparia riparia 

One observed by ALA and D. A. Turner flying over Bwawani 
swamp on 5 November 1991. Peter and Ursula Koehler (pers. comm.) 
recorded one on Chumbe island on 17 January 1993, and DBI saw a 
single bird among Eurasian Swallows Hirundo rustica over Bwawani 
swamp on 11 November 1993. 


EAST COAST AKALAT Sheppardia gunningi 

A single sight record, believed to be of this species, was made by 
H. J. Beentje in December 1989 in Jozani forest. Subsequently ALA 
netted and ringed an adult on 27 January 1991 and DBI netted and 
ringed four individuals between 13 November 1994 and 4 April 1995, 
all in Jozani forest. 


SEDGE WARBLER A crocephalus schoenobaenus 


DBI recorded three in reeds at Bwawani swamp on 2 April 1994 and 
at least six with Lesser Swamp Warblers A. gracilirostris between 17 
and 30 April 1994. 


OLIVACEOUS WARBLER Hippolais pallida 


A single bird well observed and identified by Peter and Ursula 
Koehler (pers. comm.) at Chumbe island on 13 January 1994. 


WOOD WARBLER Phylloscopus sibilatrix 
This scarce palaearctic migrant to Eastern Africa was identified by 


DBI on Chumbe island 21 March 1995; a single bird feeding in a 


Casuarina tree observed at close quarters for an hour. 


WILLOW WARBLER Phylloscopus trochilus 


Two sightings, made by DBI, of single birds in the People’s Garden, 
Stone town, on 6 and 20 April 1994. 


C. T. Collins & T. A. Araya 172 Bull. B.O.C. 1998 118(3) 


STRAW-TAILED WHYDAH V2dua fischeri 


A male was seen at Fukuchani ruins by ALA and Andrew Browning 
on 2 May 1992 with Pin-tailed Whydahs V. macroura and 
Orange-winged Pytelias Pytelia afra in the vicinity. A second sighting, 
by DBI, was made near Bwawani swamp on 6 April 1994. It is possible 
that these were escaped cage birds but the species is recorded from Dar 
es Salaam. 


EASTERN PARADISE WHYDAH V dua paradisaea 


‘Two wardens on Chumbe island watched a male at close range as it 
fed near the west cliff, in March 1994. Their detailed description was 
clearly of this species. 


Acknowledgements 


For help in the preparation of this paper, we are indebted to D. A. Turner. 


References: 

Archer, A. L. & Turner, D. A. 1993. Notes on the endemic species and some additional 
new birds occurring on Pemba Island, Tanzania. Scopus 16: 94—97. 

Britton, P. L. (ed.) 1980. Birds of East Africa. East African Natural History Society, 
Nairobi. 

Koehler, P. & Koehler, U. 1993. Notable records from Chumbe Island, Zanzibar. Scopus 
16: 115-116. 

Koehler, P. & Koehler, U. 1994. Zanzibar records of Arctic Skua Stercorarius parasiticus 
and Pomarine Skua S. pomarinus. Scopus 18: 116-117. 

Pakenham, R. H. W. 1979. The Birds of Zanzibar and Pemba. British Ornithologists’ 
Union Check List 2, Tring. 


Addresses: A. L.. Archer, P.O. Box 15676, Nairobi, Kenya. D. B. Iles, Waylands, East 
Hanney, Wantage, Oxon. OX12 OJE, U.K. 


© British Ornithologists’ Club 1998 


Natal pterylosis of tanagers II: Tachyphonus, 
Ramphocelus and Tangara 


by Charles T. Collins & Tamara A. Araya 
Received § September 1997 


Although the natal pterylosis of Neotropical passerine birds has been 
given some attention (Collins 1963, 1973; Collins & Bender 1977, 
Wimer & Collins 1994) there still remain wide gaps in our know- 
ledge. Often, generalizations about large and diverse families (i.e. 
Furnariidae) can only be made on the basis of a few specimens 
representing a small number of species and genera (Collins et al. 1991). 
Our knowledge of the natal pterylosis of tanagers (Thraupidae; 235 
species in 58 genera, Paynter & Storer 1970) is currently limited to 
studies of 6 species in 4 genera (Collins 1963, 1973, Ingels 1979, 


C. T. Collins S& T. A. Araya ‘73 Bull. B.O.C. 1998 118(3) 


Wetherbee 1958). In this paper we present data on 3 additional species 
in 3 additional genera (Tachyphonus, Ramphocelus, and Tangara) of this 
diverse assemblage of largely Neotropical birds. We also present 
supplementary data on two species of Thraupis considered earlier 
(Collins 1963, Ingels 1979). 

Counts of natal downs were made from 26 specimens examined 
under a binocular dissecting microscope. ‘The number and distribution 
of natal downs (neossoptiles) were recorded (Table 2, 3) using the 
terminology for neossoptile tracts and regions within tracts proposed 
by Wetherbee (1957) and Collins & Keane (1991). 

Six specimens of Tachyphonus rufus (all Stage A, Wetherbee 1957: 
356) were examined: two collected from each of 2 nests on 19 June 1963 
and 26 July 1964 in Arima Valley, Trinidad, and 2 from one nest on 22 
July 1976 in Los Anaucos, Est. Miranda, Venezuela. Six specimens of 
Ramphocelus carbo (4 Stage A, 2 Stage C) were examined: 3 were 
collected from one nest in 1963 (date not recorded), 2 from a nest on 22 
July 1964 and one from a nest on 22 September 1964, all from the 
vicinity of Arima or Arima Valley, Trinidad. T'wo specimens of 
Tangara mexicana were examined; both were collected from one nest on 
19 May 1964 at Waller Field, Trinidad. Three new specimens of 
Thraupis palmarum (Stage A) were examined: one collected on 6 May 
1963, and two from one nest on 3 and 4 August 1964, all in Arima 
Valley, Trinidad. Two additional specimens (Stage A) collected at the 
same site in 1962 (Collins 1963), were re-examined in this study. Seven 
specimens of Thraupis episcopus were examined: a single chick was 
collected on 18 June 1964 (Stage B), and 2 from each of three nests on 
20 June 1963 (Stage A), 18 June 1964 (Stage A), and 19 July 1964 
(Stage C) all in the vicinity of Arima, Trinidad. 

The total neossoptile counts in these tanagers, all of which build 
a typically open cup type of nest, ranged from 64 in Ramphocelus carbo 
to 238 in Thraupis palmarum (Table 1), with substantially greater 
similarity within species and genera (Tables 2 & 3). Only R. carbo 
seemed uniformly lower in total neossoptiles than the other species. 
Two specimens, including the one with the lowest total neossoptile 
count (64), were somewhat older (Stage C) when loss of some downs is 
quite likely; the total for this species (Table 1) may be artificially 
low. Even so, R. carbo has appreciably more neossoptiles than the total 
of 32 reported for Euphonia violacea (Table 1), the only closed-nest 
tanager species examined to date. The total for R. carbo is more 
similar to that of Tersina viridis (77-107 total neossoptiles; Collins 
1973), a unique fruit-eating tanager which builds its open nest 
within cavities and burrows (Schaefer 1953, Collins pers. obs.). As 
previously documented for the Tyrannidae (Collins & McDaniel 1989), 
closed-nest species typically have few or no neossoptiles while open cup 
nesting species may have 154-607 neossoptiles (Wetherbee 1957, 
Collins unpublished). The one specimen of a temperate zone tanager 
examined to date, Piranga olivacea (Wetherbee 1958), had a total of 227 
neossoptiles. This is only marginally less than the largest totals 
reported here (Table 1) for these tropical species. Intrageneric 
comparisons have shown temperate species to have more neossoptiles 


CYT. Collins S TA Avaya 174 Bull. B.O.C. 1998 118(3) 


TABLE 1 
Total neossoptile counts in tanagers (Thraupidae). n=no. of specimens 


Species Total number of neossoptiles n Source 

average (range) 
Tachyphonus rufus 132 (111-147) 6 This study 
Ramphocelus carbo 81 (64-108) 6 This study 
Tangara mexicana 160 (152-167) 2 This study 
Thraupis episcopus 164 (114-215) v) This study 
Thraupis episcopus 17. (162-175) 5 Ingels 1979 
Thraupis palmarum 209 (192-222) 3 This study 
Thraupis palmarum 175 (173-177) 2 Ingels 1979 
Thraupis palmarum 204 (184-224) 2 Collins 1963 

205 (172-238) 2 Re-examined here 

Thraupis sayaca ine! — 1 Ingels 1979 
Tersina viridis 102 (77-107) 8 Collins 1973 
Euphonia violacea 32 — 1 Collins 1963 
Piranga olivacea OOF, _— Z Wetherbee 1958 


than tropical congeners (Collins & Minsky 1982, Wimer & Collins 
1994); intergeneric comparisons, as those in this study, do not show 
this pattern as clearly. 

Partial data are available for two additional species of ‘Tangara: T. 
cayana and T. arthus (Table 2). However, these specimens were 
examined with a hand lens under field conditions and the very small 
pale neossoptiles of the primaries, secondaries and abdominal region 
may have been missed; hence the total number of neossoptiles for these 
species could not be determined. However, for those tracts and regions 
which could be examined, there is a general agreement with the 
neossoptile counts recorded for JT. mexicana (‘Table 2), suggesting an 
overall intrageneric similarity. This same problem was noted for 
thrushes which were examined in the field as part of growth studies 
(Wimer & Collins 1994). Only detailed study of preserved specimens is 
likely to produce accurate neossoptile counts. 

Two specimens of T. palmarum examined earlier (Collins 1963) were 
restudied here. The total neossoptile counts were different from the 
totals reported earlier, mostly due to documented losses of downs. 
However, in one specimen some of the very short downs on the 
primaries and secondaries were not included in the earlier counts. 
These omissions led to a higher observed total neossoptile count for one 
of the restudied specimens (‘Table 1). 

In four specimens of T. episcopus and three of T. palmarum, 1-5 
unpaired neossoptiles were noted in a medial row bordered by the 6-11 
paired downs, lateral to the midline, in the mid-dorsal region of the 
spinal tract. This is similar to the more posterior medial pelvic row, 
bordered by the lateral pelvic rows (Collins & Bender 1977), and the 
recently described interscapular region (Collins & Keane 1991), an 
unpaired medial row which overlaps the anteriormost pairs of 
mid-dorsal neossoptiles. The unique unpaired medial row noted here 


Bull. B.O.C. 1998 118(3) 


175 


C. T. Collins & T. A. Araya 


‘Qyo[/4sI1) posted sioyjO [[e ‘MOI [eIpsw poreduy y 


d d d d 981 Sst ell HE NON we li SY O6 Gel” SCs Vis CCl) ee Seed T29,. 


= = = = = = = = = = - - - (S}19A09 [eISeyed) 


- - = - — - = 1/0 1/0 (xowios [edird) 
a 0/8 0) SENS) SEO 6 eee 8 ee CC BNC eC aao 9 Li lie OR OL OPIN ee tiie 1/9 _ _(A09-"988 pry) 
8/8 —-8/8 8/8 8/6 8/6 ~—- 6/6 GORE V/s SSO Ss Sh = 8/8 6/8 L/L 6/6 6/6 6/6  (S}49A09 AIvpuOdas 1903v919) 
é é é é Wi = Cie = Gt Oe Ae = ape OCs Gt TIC SGiGe cet (saliepuosas) 
= — — - - - = — = — - - - - — - = - (s}19A09 Areutid 1938913) 
é é é é 9/¢ g/€ Lie = ViEOVe. Die = age = = = = ~ (sorsetutid) 
IvlV 
9/9 9/9 s/9 9/9 9/9 9/9 9/9 s/s 9/9 1/0 0/0 0/0 O/9 G/Oy S/S S/S — 9/9 9/9 jepneD 
= 3 = = 3 = Site ze = se oe ee. = a Ge = jeaniy 
- - - - — — = - = - - _ - - - - - _ (rey [Exe) 
é é é é yey ail ae = SG ie (OPS ue IOS = Lira (jpuropgqe) 
[eu A 
tly v/S 8/6 8/8 Wh ols 0/9 +t/p S/r 8/6 9/9 s/9 G/Se “2/8 °O/S “BIR SP G/ee “L/6 [R1OW9 yf 
9/9 = 8/L 6/8 —L/8 6/6 6/6 B/G OG TS 0) rig eo) 9 Lite Be Lie TO ST O76 qepndeog 
- = - = = — = = = - - = - - _ - ~ — y(jeurds) 
W/1 €/€ = = 1/1 1/1 WO 0/0) 0) ae 00a) eee Gg Be 0/0: ae re = (otajad ye193e]) 
Ol +1 8 8 ral Ol Lee 8 OL eee! 6 fl Ol oy OE 0 L ¥(olqed) 
ae ae 4/9 —s/9 6/01 8/8 9/9 s/s 9/s ze s/9 8/9 Oe RL Wee OPO: LIL BHO WLI 9 (]es10p-prw) 
jeuidg 
g/g (8/9 Pts Toys Sip 9/S C/O Ok me Ce NG C/N eG G/S= <OGe 0/0 OWL SYEAS0/S ({eatdi990) 
Wey *Cvial 01/6 01/8 LL Ly 4/9 8/8 O01 6/6 8/L 6/11 4/8 OVO 8/8 6/01 11/6 6/11 ({RU0109) 
jeadey 
SNYJAD DUDADI DUDIIXIU OgAv9 snjar0yTunyy snfna snuoydayav J, (UOIBO1) JOR, 
DADBUD J, DADBUD J, DADBUD J, 


sioseur) vavsuvny, pur ‘snjavoyduvy ‘snuoyddaysvy, ut yuosaid sa[iydossoaon 
Gal ley Is 


176 Bull. B.O.C. 1998 118(3) 


C. T. Collins & T. A. Araya 


‘(QJe]/1y4st41) posted s1oyjo [[e ‘Mor [eIpow poseduyy 


ELI SEZ (SOG 761 C17 971 +91 +11 SIZ S07 861 vl [e310], 
= ¢/Z = = = = = = = = = = (S119A09 [eIdejed) 
= = 1/1 = 0/1 = - 1/1 = 1/T = = (xouso1 [edie9) 
9/9 8/8 8/8 L/8 9/9 9/9 S/€ 6/L 8/8 8/8 8/8 /€ — (S}4#9A00 ArBpuodas a[ppru) 
9/6 01/01 6/6 6/6 6/6 8/8 9/8 s/€  OL/OL 6/6 6/6 €/@  (S¥49A09 Arvpuodses 103k913) 
= ye 8/8 = = 0/7 Go 7/7 c/¢ c/€ O/T 8/8 (Ssolizpuoseas) 
= = IW = oo = = = = = = = (s}19A00 AteutId 10}¥0138) 
= O1/O1 S/S ae 4/7 ~ = 0/+ 6/6 ¢/9 0/7 6/6 (sorreuttid) 
ItlV 
9/9 9/9 9/9 9/9 9/9 ¢/Z +/7 ¢/9 9/S 9/9 9/9 9/9 jepnes 
0/1 €/+ c/¢ = 4/7 = = _ AG - = = jernig 
0/1 Ge = = a = = = - = = a (1e][Ixe) 
Wl MISE OGL 6 SUS 1/2 S/+ 0/0 O/E€ 1/€ 0/0 0/0 (jeuruopqe) 
[PTUI A 
Gia sche SOG CL © C/O WO iO OT YS On eh Oye [Brows J 
01/01 6/01  Ot/Ol 6/6 Gio 8/8 Cy (0 ae Sem Ol 6/6 6/6 9/S Jepndeag 
i G 0 I 0 0 C 0 I ¢ S 0 y(Teurds) 
0/T 1/€ 1/1 +/€ Z/€ 9/S 6/€ 0/0 b/¥ L/9 S/L 0/0 (sted [e193e7) 
(GA) iat Il OI ZI 8 8 6 el Ol IT 8 x (9tAjod) 
9/S 9/L 6/L GON ICA LI€ ¢/9 9/9 6/6 TO) = 01/6 9/9 CTPSROREEE) 
jeutds 
+/¥ 1/0 ¢/S c/s ¢/S S/+ L/8 ¢/¢ B/L 8/6 11/6 +/¢ (Tetdt990) 
NE = IEE = GIO ONS oer | 6/L oV ie S=8)S ei Ce . MYC) ==F1/Siae es ([BUO0109) 
jeadea 
unsvujod sidnvay J, sndoasida sidnoay J, (UOIB91) JOR], 


Siosvur} sidnvsy J, Ut JUaseid sopIydossooNy 
€ A 1aViL 


C. T. Collins S& T. A. Araya 177 Bull. B.O.C. 1998 118(3) 


appears to represent an additional undescribed region of the spinal tract 
which we propose to call the spinal region. However, examination of 
additional species, in this or other passerine families, may show this 
spinal region to be at times contiguous with either the interscapular or 
midpelvic region and not the distinct region it appears here. The 
previous studies of Thraupis tanagers (Collins 1963, Ingels 1979) did 
not differentiate any regions within the spinal tract. 

Current evidence indicates that most neossoptiles are fully developed 
at hatching (Wetherbee 1957: 356). However, as noted by Ilyashenko 
(1984), in a few cases some neossoptiles only appear 2-3 days 
post-hatching. In one specimen of Thraupis palmarum 7 primary covert 
and 8 secondary neossoptiles were present only as largely unerupted 
dark streaks, 1-2 mm long, under the skin. The tips of 2—3 of these 
protruded for less than 1 mm through the skin; all would have fully 
erupted within a day or two of hatching. At this stage no contour pin 
feathers were observable so these streaks were clearly unerupted 
neossoptiles and not incoming teleoptiles. Such unerupted neossoptiles 
should not be equated with the dense coat of whitish “secondary down’’ 
which erupts 5-7 after hatching in Elaenia flycatchers (Skutch 1960: 
303). These “‘secondary downs” are located in “regions between the 
tracts of natal down’”’ (Skutch 1960). They are not associated with 
incoming teleoptiles and thus cannot be true neossoptiles. They are 
probably early-appearing down or semiplume portions of the juvenal 
(teleoptile) plumage as previously reported for Notiochelidon cyanoleuca 
(Hirundinidae; Arnold et al. 1983). Other reports of second coats of 
natal down in passerines (Erithacus, Lucinia; Cramp 1988) should be 
critically reexamined. 

The great diversity in latitudinal range and nesting habits of the 
tanagers makes them an appropriate group in which to examine in 
more detail some of the ecological correlations previously noted in 
other families. However, as pointed out earlier (Collins 1990), more 
specimen material will have to be collected before such studies are 
possible. 


Acknowledgements 


Field work in Trinidad (C. T. C.) was supported by grants from the Frank M. Chapman 
Memorial Fund of the American Museum of Natural History. The study of these 
specimens (T. A.) was part of the Young Scholars Program of the National Science 
Foundation (ESI-9255941) to W. C. Ritz. Kristin Bender assisted with the study of some 
specimens reported here. 


References: 

Arnold, K. A., Boyd, E. J. & Collins, C. T. 1983. Natal and juvenal plumages of the 
Blue-and-white Swallow (Notiochelidon cyanoleuca). Auk 100: 203-205. 

Collins, C. T. 1963. Natal pterylosis of tanagers. Bird-Banding 34: 36-38. 

Collins, C. T. 1973. The natal pterylosis of the Swallow-tanager. Bull. Brit. Orn. Cl. 93: 
155-157. 

Collins, C. T. 1990. Intraspecific variation in the natal pterylosis of the Ochre-bellied 
Flycatcher Myionectes oleagineus (Tyrannidae). Bull. Brit. Orn. Cl. 110: 143-145. 

Collins, C. T. & Bender, K. E. 1977. Cervical neossoptiles in a Neotropical passerine. 
Bull. Brit. Orn. Cl. 97: 133-135. 


D.L. Anderson, M. Bonta & P. Thorn 178 Bull. B.O.C. 1998 118(3) 


Collins, C. T. & Keane, K. 1991. Natal pterylosis of phoebes. Wilson Bull. 103: 300-303. 

Collins, C. T., Marin A., M. & Lentino R., M. 1991. Natal pterylosis of Premnoplex 
brunnescens, Thripadectes virgaticeps and Synallaxis candei (Furnariidae). Bull. Brit. 
Orn. Cl. 99: 12-15. 
Collins, C. T. & McDaniel, K. M. 1989. The natal pterylosis of closed-nest building 
tyrant flycatchers (Aves: Tyrannidae). Bull. So. Calif. Acad. Sci. 88: 127-130. 
Collins, C. T. & Minsky, D. 1982. Natal pterylosis of three Neotropical blackbirds 
(Icteridae). Bull. Brit. Orn. Cl. 102: 129-131. 

Cramp, S. (ed.) 1988. The Birds of the Western Palearctic. Vol. 5. Tyrant Flycatchers to 
Thrushes. Oxford Univ. Press. 

Ilyashenko, V. Y. 1984. A new downy pteryla in passerine birds. Bull. Brit. Orn. Cl. 104: 
53-54. 

Ingels, J. 1979. Natal pterylosis of three Thraupis tanagers. Bull. Brit. Orn. Cl. 99: 12-15. 

Paynter, R. A., Jr. & Storer, R. W. 1970. Check-list of Birds of the World. Vol. 13. Mus. 
Comp. Zool., Cambridge, MA. U.S.A. 

Schaefer, E. 1953. Contribution to the life history of the Swallow Tanager. Auk 70: 
403-460. 

Skutch, A. 1960. Life histories of Central American birds. Pacific Coast Avifauna 34: 
1-593. 

Wetherbee, D. K. 1957. Natal plumages and downy pteryloses of passerine birds of 
North America. Bull. Am. Mus. Nat. Hist. 113: 339-436. 

Wetherbee, D. K. 1958. New descriptions of natal pterylosis of various bird species. 
Bird-Banding 29: 232-236. 

Wimer, M. C. & Collins, C. T. 1994. Natal pterylosis of some neotropical thrushes 
(Muscicapidae: Turdinae). Bull. Brit. Orn. Cl. 114: 170-173. 


Addresses: Dr. C. T. Collins, Department of Biological Sciences, California State 
University, Long Beach, California 90840, U.S.A. Ms. T. A. Araya, Long Beach 
Polytechnic High School, 1600 Atlantic Avenue, Long Beach, California 90813, 
UsS:A: 


© British Ornithologists’ Club 1998 


New and noteworthy bird records from 
Honduras 


by David L. Anderson, Mark Bonta 5 Pilar Thorn 
Received 23 September 1997 


From an ornithological standpoint, Honduras remains one of the least 
studied countries in Central America. Little has been published since 
Monroe’s survey in 1968. This paper is based on over 25 cumulative 
years of residence in Honduras (DA 3.5, MB 3.5, PT 20), during which 
the observation of birds was a central aspect of our work. We report 
here significant information on 17 species, including 8 previously 
unreported for Honduras. We also update information for some species 
already recorded in Honduras, but whose status is poorly substantiated 
in the literature or which have not been reported since Monroe. 

The majority of our sightings originated from fieldwork in two main 
areas: Las Marias, in the Rio Platano watershed 25 km south of the 
Caribbean coast, Dpto. Gracias a Dios; and the Sierra de Agalta 
mountain range and surrounding communities, principally Gualaco, 


D.L. Anderson, M. Bonta © P. Thorn 179 Bull. B.O.C. 1998 118(3) 


E] Boqueron, and La Venta, in Dpto. Olancho. Las Marias is situated 
in the Rio Platano Biosphere Reserve of the Honduran Mosquitia 
region in the midst of more than 250,000 ha of lowland rain forest with 
few human inhabitants and scant disturbance. The Sierra de Agalta 
retains mixed pine-oak woodlands, cloud forests, and lower montane 
wet forests, with the surrounding valleys dominated by agriculture. It 
is contiguous with the lowland forests of the Mosquitia. 


WHITE PELICAN Pelecanus erythrorhynchos 

Although not previously reported for Honduras, PT has observed 
this species on both the Pacific and Caribbean coasts. In March 1993 
three individuals were observed with eight Brown Pelicans as they 
rested on a beach of Laguna de Los Micos, 10 km west of Tela, Dpto. 
Atlantida. Local residents told her that these birds had been appearing 
at the same beach for the last 2—3 years. PT first heard reports of White 
Pelicans in the Bay of Fonseca in 1993 and on 28 June 1997 personally 
observed more than 30 near San Bernardo, Choluteca Department. 


SEMIPLUMBEOUS HAWK Leucopternis semiplumbea 


This species was previously reported only twice in Honduras, and 
but once this century (Monroe 1968, Marcus 1983). DA made 
observations of 6 different individuals of this species from 25 January 
to 27 May 1996 while working in the vicinity of Las Marias. One 
observation (8 March) included a prey transfer of an unknown lizard 
species to a bird that may have been immature, based on faint white 
streaking on the face. Five of the sightings were in primary, lowland 
rain forest, and the remaining one in regenerating forest surrounded by 
primary forest and agricultural plots. They were distinguished from the 
similar Leptodon cayenensis, which was also present locally, by their 
shorter tail, yellow eye, orange cere and legs, and smaller size. 


HARRIS’ HAWK Parabuteo unicinctus 


DA observed one adult at close range on 16 January 1993 near 
Monjaras, Dpto. Choluteca. The bird was flying 6m high over arid 
thorn scrub habitat. PT has also seen individuals in this vicinity. The 
sooty brown plumage with rufous thighs and wing coverts made the 
birds unmistakable. This species has not been previously reported from 
Honduras. The nearest published observations were in Dpto. La Paz 
and Usulatan, in El] Salvador (Thurber et al. 1987). 


HARPY EAGLE Harpia harpyja 


DA photographed a Harpy Eagle on 12 April 1996, 12 km southwest 
of Las Marias at a site known as Walpulbantara Petroglyphs along the 
Rio Platano. The bird was observed for 24 min, and appeared to be 
perch hunting (flying from tree to tree and actively scanning the 
vicinity). Massive yellow tarsi, fine black barring on the thighs, white 
underparts, black breast, and gray double crest clearly separated this 
species from Morphnus guianensis. he scapulars were an ashy gray, 
indicating the bird was not a full adult. The habitat was primary rain 
forest. Additionally, two adults were photographed together in the 


D.L. Anderson, M. Bonta & P. Thorn 180 Bull. B.O.C. 1998 118(3) 


Valle Sutawala, Dpto. Gracias a Dios in October 1991 (V. Murphy 
pers. comm.). ‘Together these observations suggest a _ breeding 
population in Honduras. 


BLACK-AND-WHITE HAWE-EAGLE Spizastur melanoleucus 

Other than a decapitated head of this species reported by Howell & 
Webb (1992) in Tela, Dpto. Atlantida, there have been no published 
records since 1950 (Monroe 1968). DA observed an adult flying over 
Cerro Ubhjuri, 4km northeast of Las Marias on 2 April 1996, and 
another adult both perched and flying in the Quebrada Wahawala 
drainage 6 km southwest of Las Marias on 1 and 3 April 1997. They 
were distinguished from other white raptors by the presence of a red 
cere, black lores, small black crest, completely white underparts 
including the tarsi, and blackish upperparts with white wing coverts. 
Both birds were observed in primary rain forest. 


WHITE-THROATED CRAKE Laterallus albigularis 


Monroe (1968) only knew of one record for this species in Honduras 
and its range in relation to that of the Ruddy Crake L. ruber is poorly 
known (Howell & Webb 1995). DA found this species to be ubiquitous 
in tall grasses along the Rio Platano. ‘Two separate family groups (one 
with a single chick and one with two) were observed on 25 April 1997 
and the long trilling calls were heard throughout the months 
January-June. No calls of L. ruber, with which DA is also familiar, 
were heard. 


BLACK-AND-WHITE OWL Ciccaba nigrolineata 

‘There have been only two reports for this species in Honduras 
including a specimen collected in 1891 (Monroe 1968, Howell & Webb 
1992). Both observations were from the Caribbean coast. We found 
another specimen preserved as a live mount in the collection of the 
Universidad Nacional Autonoma de Honduras in ‘Tegucigalpa. ‘The 
bird was collected in October 1979 near San Marcos de Colon, Dpto. 
Choluteca, in southern Honduras. San Marcos is the site of some 
still intact, but rapidly dwindling, dry forests. MB also observed 
a Black-and-white Owl at close range in the Sierra de Agalta on 
2 February 1996. This species is apparently well-distributed in the 
country, and the paucity of sightings most likely the result of its 
nocturnal habits. 


STRIPED OWL Aszo0 clamator 

This species is apparently rare north of Panama and was previously 
known from Honduras by a single specimen (Monroe 1968). DA 
skinned a second bird collected in 1993 in the vicinity of Comayagua in 
Dpto. Comayagua; the specimen was deposited in the Louisana State 
University Museum of Natural Science collection (LSUMZ# 161,378). 
The bird had died after being captured as a nestling in grassy swales 
near the city. 


OCELLATED POORWILL Nyctiphrynus ocellatus 


There is only one confirmed record of this species north of Costa 
Rica (Eisenmann 1955), and in Costa Rica it is known from a single 


D. L. Anderson, M. Bonta & P. Thorn 181 Bull. B.O.C. 1998 118(3) 


breeding population near Brasilia (Stiles & Skutch 1989). DA found a 
female incubating two eggs on 16 May 1997, 2 km southwest of Las 
Marias (photographs deposited in the WF VZ and LSUMZ collections) 
and heard the calls of this species frequently at night in 1996 and 1997. 


BRONZY HERMIT Glaucis aenea 

This species is one of the most common hummingbirds in Heliconia 
thickets in the Rio Platano watershed. On 16 February 1996 an 
individual flew into a window in Las Marias. When caught and released 
it dropped several tail feathers which have been deposited in the 
Western Foundation of Vertebrate Zoology (WF VZ, 53-181). This is 
the first report of this species for Honduras. 


HONDURAN EMERALD Amazilia lucia 


On 2 January 1996, MB confirmed the existence of a population of 
this species 30 km northeast of Gualaco in the Agalta Valley. Upon 
return visits by MB and DA (31 January 1996, 21 June 1997) it was the 
most common hummingbird at this locality. The available habitat of 
thorn forest consists of less than 100 ha, all on private land, at an 
elevation of 500 m above sea level. This forest is rapidly being cleared 
for conversion to rice fields and cattle pastures. MB has searched 
extensively in the Guayape Valley where the species was collected in 
1937 (Monroe 1968) but was unable to locate any remaining suitable 
habitat. Conservation programmes for this species are urgently needed. 
This record establishes only the second known remaining area of 
occurrence (combining “‘Coyoles” and ‘“‘Olanchito’’; Howell & Webb 


1989). 


GREEN-AND-RUFOUS KINGFISHER Chiloroceryle inda 


On 2 June 1997 DA observed a male of this species as it flew up a 
densely wooded stream 2 km northwest of Las Marias and perched 
nearby for 5 min. The bird was distinguishable by its glittering green 
back, buff throat, and deep rufus breast and undertail coverts. It was 
clearly larger than the Pygmy Kingfisher Chloroceryle aenea, also 
present in the area, and lacked white on its underparts. This is the first 
record of this species for Honduras. 


FULVOUS-BELLIED ANTPITTA Hylopezus fulviventris 

Although recently considered of doubtful occurrence in Honduras 
(Howell & Webb 1995), DA found this species to be common in 
successional habitats along the Platano River, especially where land 
cleared for agriculture has been left fallow and allowed to regenerate 
into dense thickets. Three males with enlarged testes were collected 
while singing and have been deposited in the WFVZ collection 
(52-988—-52-990). This represents a 450 km northward extension of the 
known range of this species (Monroe 1968). 


THREE-WATTLED BELLBIRD Procnias tricarunculata 


Although previously reported from the Sierra de Agalta (Monroe 
1968), recent accounts of this species state that it does not breed there 


D. L. Anderson, M. Bonta & P. Thorn 182 Bull. B.O.C. 1998 118(3) 
(Howell & Webb 1995, Wege & Long 1995). We have found calling 


males exceedingly common in May and June in montane wet forests 
from 600 to 2,100 m elevation, and have recorded them in most other 
months of the year (March, April, July, August, November). We have 
also frequently heard the raspy, wavering calls of birds we suspect to be 
immature males. We feel that this information strongly indicates the 
presence of a breeding population in Honduras. 


WHITE-VENTED EUPHONIA Euphonia minuta 


MB made repeated observations of a male in January 1996 in La 
Venta, Dpto. Olancho in bottomland habitats along the Rio Mataderos. 
‘The bird was seen feeding in mixed flocks of other Euphonia species (E. 
affins and E. hirundinacea) and was identified by its clearly white 
undertail coverts. This is the first record for Honduras, although the 
species has been recorded from Guatemala and Nicaragua. 


NORTHERN CARDINAL Cardinalis cardinalis 


P’'T first discovered Northern Cardinals on the Swan Islands, 200 km 
north of the Honduran mainland, in June 1996 when 2-3 males were 
observed singing. The species has been seen there in subsequent visits, 
with 4-10 birds observed on any given visit. Given the distance from 
the nearest known breeding population (Belize) and the isolation of the 
islands from the mainland, we suspect that these birds are escapees, a 
view corroborated by local residents. We are unsure of the current 
population trend. This is the first report of this species for Honduras. 


WHITE-EARED GROUND-SPARROW MVlelozone leucotis 


MB has made repeated observations of White-eared Ground 
Sparrows at 1150 m on Cerro Agua Buena in the El Boqueron National 
Monument, Dpto. Olancho since 11 February 1993. An adult was 
flushed from a nest containing one egg and one nestling on 27 May 
1997 in the same vicinity, and one adult and a nest were photographed 
on 31 July 1997 (photographs deposited in LSUMZ). The birds are 
typically found in the thick undergrowth of 7 m tall regenerating forest. 
The species is known from the highlands of Chiapas, Guatemala, El 
Salvador, and Nicaragua, but this is the first record for Honduras 
(Ridgely & Gwynne 1989). 


Acknowledgements 


Permits to carry out scientific studies and make biological collections were granted by 
COHDEFOR, Republic of Honduras. L. Kiff provided constructive criticisms of an 
earlier version of the manuscript. V. Remsen, R. Ridgely, R. Colorado, and C. Sumida 
assisted with specimen curation and identification. 


References 

Eisenmann, W. 1955. The species of Middle American birds. Trans. Linn. Soc. New 
York ail 

Howell, S. N. G. & Webb, S. 1989. Notes on the Honduran eed: Wilson Bull. 101: 
642 643. 


B. L. Lanis, R. P. Clay & F.C. Lowen 183 Bull. B.O.C. 1998 118(3) 


Howell, S. N. G. & Webb, S. 1992. New and noteworthy bird records from Guatemala 
and Honduras. Bull. Brit. Orn. Cl. 112: 42-49. 

Howell, S. N. G. & Webb, S. 1995. Birds of Mexico and Northern Central America. 
Oxford University Press. 

Marcus, M. J. 1983. Additions to the avifauna of Honduras. Auk 100: 621-629. 

Monroe, B. L., Jr. 1968. A distributional survey of the birds of Honduras. Orn. Monogr. 
no. 7, A.O.U. 

Ridgely, R. S. & Gwynne, J. A. 1989. A guide to the birds of Panama, 2nd edn. Princeton 
Univ. Press. 

Stiles, R. G. & Skutch, A. F. 1989. A guide to the birds of Costa Rica. Cornell Univ. Press. 

Thurber, W. A., Serrano, J. F., Sermeno, A. & Benitez, M. 1987. Status-of uncommon 
and previously unreported birds of El Salvador. Proc. Western Found. Vert. Zool. 3: 
109-293. 

Wege, D. C. & Long, A. J. 1995. Key areas for threatened birds in the neotropics. 
BirdLife Conservation Series no. 5. Burlington Press, Cambridge. 


Address: David L. Anderson, Department of Biology, Boise State University, 1910 
University Drive, Boise, Idaho 83725, U.S.A. Mark Bonta, Department of 
Geography and Anthropology, Louisiana State University, Baton Rouge, LA 70803, 
U.S.A. Pilar Thorn, Departamento de Biologia, Universidad Nacional Autonoma de 
Honduras, Tegucigalpa, Honduras, Central America 


© British Ornithologists’ Club 1998 


A new plumage of the White-winged Nightjar 
Caprimulgus candicans (Aves: Caprimulgidae) 


by Bernabé Lopez Lanus, Robert P. Clay & 
James C. Lowen 


Received 30 September 1997 


The White-winged Nightjar Caprimulgus candicans is one of the least 
known caprimulgids in the Americas (Collar et al., 1992, Lowen et al., 
1996b) and is considered Critically Endangered (Collar et al., 1994). It 
is known from two specimens collected in the cerrado of central Brazil: 
one at Irisinga, Sdo Paulo, on 6 January 1823 (not February as in Collar 
et al., 1992), and the other at Cuyoba, Mato Grosso, in either 
December 1823 or June 1825 (both dates on label; N. Cleere zn litt. 
1997). There is older evidence from Paraguay (de Azara 1802-05), but 
without details of date or locality. Subsequent records have come from 
only three widely spaced sites: Emas National Park, Goias state, Brazil 
(Collar et al., 1992), Beni Biological Station, Yucuma province, Bolivia 
(Davis & Flores 1994), and Reserva Natural del Bosque Mbaracayu 
(RNBM), Canindeyu department, Paraguay (Lowen et al., 1996b, 
1997). 

During fieldwork at RNBM in September and December 1995, we 
(principally BLL) captured six male C. candicans in presumed 
definitive plumage and a seventh C. candicans of a_ previously 
undescribed plumage. This seventh individual lacked any white in the 
wings and tail, but on the basis of structure, head pattern and overall 


BIL Lani RS PACTayVXS F.C. Lowen ° 184 Bull. B.O.C. 1998 118(3) 


plumage tones, was clearly C. candicans. In addition, the biometrics of 
this bird fall within the range of that now known for C. candicans (see 
Appendix). The bird was caught in a small area of campo 
sucio—grassland with scattered shrubs and trees, the most common 
being dwarf palms Butia paraguayensis—with many anthills. All male 
C. candicans in definitive plumage caught and seen frequented this 
same area. No other caprimulgids were observed here. 

‘The Paraguay observations revealed the extent of white in the wing 
of male C. candicans to be variable. Variation in the wing and tail 
patches of nightjars has been rarely considered (Forero & Tella 1997; 
although see Cramp 1985, Ingels & Ribot 1982, Jackson 1984), but, for 
one species (Red-necked Nightjar Caprimulgus ruficollis), has been 
shown to be age-related (Forero et al., 1995). Male C. candicans 
exhibiting the maximum extent of white in the wing were assumed to 
be in definitive plumage. 

Such males have blackish primary tips; white primary bases, 
secondaries, primary coverts and greater coverts (all fringed blackish); 
wholly white underwing except for blackish primary tips; white 
undertail and uppertail save for the central pair of rectrices; and white 
lower breast and belly. The species has a distinctive head pattern with 
pale greyish-brown forehead, crown and nape very finely barred and 
vermiculated dark brown. The central crown feathers are dark brown 
to blackish, barred with tawny, whilst the extreme sides of the crown 
are cinnamon finely barred brown. Extending from the base of the bill 
above the eye and just beyond is a fine whitish stripe, bordered on 
either side by cinnamon. The lores and ear coverts are dark chestnut 
barred blackish, forming a distinct dark eye patch. A distinctive 
feature is a broad whitish submoustachial stripe. In-hand photographs 
of a male in definitive plumage were published in Lowen et al. (1996a) 
and ‘Tobias & Lowen (1996), with a field photograph in Lowen 
(1997). 


Description of the new plumage 


The seventh (non-definitive male) individual was captured by BLL on 
8 December 1995. It was considered unwise to collect the bird given 
uncertainty as to the species’ local and global status. Feather samples 
were taken and have been deposited at the Museo Nacional de Historia 
Natural del Paraguay (Asuncion). A full description was taken at the 
time of capture, and feather samples were subsequently compared to 
a table of colours (Smithe 1975). Colours mentioned in uppercase 
(e.g. ““Burnt Umber’’) refer to feather samples compared directly to 
Smithe; colours in lowercase (e.g. ““‘blackish’’) pertain to the original 
description. A description of the bird is as follows: 

Bare part coloration as the captured males in definitive plumage: iris 
chestnut; periocular membrane cinnamon; upper and lower mandibles 
blackish with their bases flesh coloured; nostril protruberances also 
flesh coloured; legs and feet flesh with a grey tone; nails blackish. 

Head pattern also much as the males captured, with centre of crown 
Burnt Umber, sides of crown greyish with indistinct fine black and 


B. L. Lanis, R. P. Clay & F.C. Lowen 185 Bull. B.O.C. 1998 118(3) 


white markings. Ocular region dark cinnamon-rufous with whitish 
submoustachial line. Hindneck, mantle, back and wing-coverts Light 
Clay to Cinnamon with Drab fine spotting, barring and vermiculations. 
Base of scapular feathers similar, but towards tip Cinnamon with 
Dusky Brown spots along shaft and small white apical spot. Outermost 
primary Dark Greyish Brown. Second primary (numbered ascend- 
antly) Dark Greyish Brown with six equally spaced Cinnamon spots on 
each web, forming transverse bars, a pattern repeated over the 
remaining primaries and all the secondaries and tertials. The four 
outermost primaries with inconspicuous small white terminal spots. 
Prominent emarginations to second, third and fourth primaries. 
Central rectrices Drab and with slight Dark Raw Umber bars. Outer 
rectrices with equally spaced Dusky Brown and Cinnamon bars. 
Undertail similar in pattern but paler. Throat dark cinnamon-rufous 
contrasting with the rest of the underparts. Foreneck and flanks 
Cinnamon with fine Dark Greyish Brown bars. Breast similar but with 
feathers tipped white, thus forming small whitish spots. Belly, vent and 
undertail coverts white. Thighs light cinnamon contrasting with the 
rest of the lower underparts. 

The bird showed signs of neither moult nor feather wear, appearing 
to be in very fresh plumage. Given that it was caught at what is likely to 
be towards the end of the breeding season in Paraguay (Lopez Lants 
et al. in prep.), the bird would thus appear to be an immature. The 
exact shape of the outer primaries and rectrices of caprimulgids is a 
reliable indicator of their age, with adults exhibiting broad feathers 
with rounded tips whilst those of immatures are narrow and pointed, 
often with pale tips to the primaries (N. Cleere zm litt. 1996, RPC pers. 
obs.). Sample outer rectrices and primaries were relatively narrow and 
pointed, indicating the bird to be an immature. The small white 
terminal spots to the outer four primaries (a feature not shown by males 
in definitive plumage) support this diagnosis. 

The all dark outer primary (albeit with terminal white spot), 
contrasting with barred remaining primaries, is intriguing, since the 
remiges of immature Caprimulgids are generally uniform prior to 
moult (Cleere in press). However, since Caprimulgids (including the 
type specimen of C. candicans) moult primaries descendantly (Cleere in 
press), it seems unlikely that the bird was in moult, exhibiting new, 
adult-type outermost primaries among otherwise immature remiges on 
both wings. The all dark outer primary of this bird thus appears to be 
typical of immature male C. candicans. 'The outer primary of males in 
definitive plumage is noticeably modified, apparently to aid production 
of the mechanical sounds that accompany display (Lopez Lanus et al. 
in prep.). In this context, it is less surprising that the outer primary of 
immature males differs so markedly. 

The plumage of adult female C. candicans has yet to be formally 
described, but apparent females have been discovered during 1997 
fieldwork at RNBM (J. Mazar Barnett, D. Capper & R. Pople verbally 
1997, RPC pers. obs.). Neither these birds, nor the immature collected 
in the mid 1820s (sex unknown, but possibly female; Cleere in press), 
show white underparts, and cinnamon bands extend across all the 


By) Wants, RYUPNClay\S F.C. Lowen Ws Bull. B.O.C. 1998 118(3) 


primaries in both (J. Mazar Barnett, D. Capper & R. Pople verbally 
1997, RPC pers. obs., Cleere in press). ‘The largely white underparts 
and dark outer primary—characteristics common to males in definitive 
plumage and to the 1995 individual described here—would indicate 
this new plumage to be that of an immature male. 


Conclusions 


On the basis of head pattern, overall plumage tones, bare part 
coloration, structure and biometrics, this plumage clearly corresponds 
to C. candicans. Several factors allow the diagnosis of this plumage as 
that of an immature bird, perhaps most likely an immature male. 
Surveys in what remains of the optimal habitat of the species 
(apparently relatively open campo sucio) throughout its range are 
urgently needed to assess its true status. 


Acknowledgements 


Fieldwork was conducted under the banner of “‘Project Yacutinga ’95”’, for which full 
acknowledgements are given in Lowen et al. (1996b). We thank our colleagues during the 
project, particularly D. Liley, J. Mazar Barnett, M. Pearman and J. Tobias. In Paraguay 
we thank the staff of the Fundacion Moisés Bertoni, particularly L. Bartrina, E. Esquivel, 
A. Madrono N., A. Yanosky and the guards at RNBM. In Argentina, BLL would like to 
thank the Asociacion Ornitologica del Plata and R. Straneck for their help, and J. 
Leiberman for photographs of the feather samples. An earlier version of this note 
benefited from the comments of N. Cleere, N. Collar, C. Feare, R. Fraga, D. Wege and 
an anonymous reviewer. N. Cleere provided details of the two nineteenth-century 
specimens and the draft text from his forthcoming family monograph. 


References: 

de Azara, F. 1802-05. Apuntamientos para la Historia Natural de los Pajaros del Paraguay 
y Ruio de la Plata. Vol. 1 and 2. Imprenta de la Viuda de Ibarra, Madrid. 

Cleere, N. 1998. Nightjars Pica Press, Sussex. 

Collar, N. J., Crosby, M. J. & Stattersfield, A. J. 1994. Birds to Watch 2: the world list of 
threatened birds. BirdLife International, Cambridge. 

Collar, N. J., Gonzaga, L. P., Krabbe, N., Madrono Nieto, A., Naranjo, L. G., Parker, 
T. A. & Wege, D. C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red 
Data Book (Third Edition, Part 2). International Council for Bird Preservation, 
Cambridge. 

Cramp, S., ed. 1985. The birds of the western Palearctic, Vol. 4. Oxford University Press, 
Oxford. 

Davis, S. & Flores, E. 1994. First record of White-winged Nightjar Caprimulgus 
candicans for Bolivia. Bull. Brit. Orn. Cl. 114: 128-129. 

Forero, M. G., Tella, J. L. & Garcia, L. 1995. Age-related evolution of sexual 
dimorphism in the Red-necked Nightjar (Caprimulgus ruficollis). ~. Orn. 136: 
447-451. 

Forero, M. G. & Tella, J. L. 1997. Sexual dimorphism, plumage variation and species 
determination in nightjars: the need for further examination of the Nechisar Nightjar 
Caprimulgus solala. Ibis 139: 407-409. 

Hayes, F. E. 1995. Status, distribution and biogeography of the birds of Paraguay. 
American Birding Association: Monog. Field Orn. 1. 

Ingels, J. & Ribot, J. H. 1982. Variations in the white markings of the Blackish Nightjar 
Caprimulgus nigrescens. Bull. Brit. Orn. Cl. 102: 119-122. 

Jackson, H. D. 1984. Key to the nightjar species of Africa and its islands (Aves: 
Caprimulgidae). Smithersia 4: 1-5. 

Lopez Lanis, B.,. Clay, R: P.,.lowen, J. C.,° Mazar, Barnett, |), Wadsontg. sess 
Pearman, M. & Tobias, J. A. In prep. The display of the White-winged Nightjar 
Caprimulgus candicans with notes on its ecology. 


In Brief | 187 Bull. B.O.C. 1998 118(3) 


Lowen, J. C. 1997. Projects Canopy ’92 and Yacutinga ’95. World Birdwatch 19: 18. 

Lowen, J. C., Bartrina, L., Brooks, T. M., Clay, R. P. & Tobias, J. A. 1996a. Project 
YACUTINGA ’95: Bird surveys and conservation priorities in eastern Paraguay. 
Cotinga 5: 14-19. 

Lowen, J. C., Bartrina, L., Clay, R. P. & Tobias, J. A. 1996b. Biological surveys and 
conservation priorities in eastern Paraguay. CSB Conservation Publications, 
Cambridge. 

Lowen, J. C., Clay, R. P., Mazar Barnett, J.. Madrono N., A., Pearman, M., Lopez 
Lanus, B., Tobias, J. A., Liley, D. C., Esquivel, E. Z., Reid, J. M. & Brooks, T. M. 
1997. New and noteworthy observations on the avifauna of Paraguay. Bull. Brit. 
Orn. Cl. 117: 275-293. 

Smithe, F. B. 1975. Naturalist’s Color Guide. The American Museum of Natural History, 
New York. 

Tobias, J. & Lowen, J. 1996. Strange-tailed Tyrants and White-winged Nightjars. 
Birding World 9: 361-363. 


Address: B. Lopez Lanus, Asociacién Ornitologica del Plata, 25 de Mayo 749., Piso 2 
*“6’’, (1002) Buenos Aires, Argentina; R. P. Clay, Large Animal Research Group, 
Department of Zoology, University of Cambridge, Downing Street, Cambridge 
CD2 3EJ. U.K.; J. C. Lowen, BirdLife International, Wellbrook Court, Girton 
Road, Cambridge CB30NA, U.K. (J.C.L. present address: Department of 
Environment, Transport and the Regions, Rose Court, 2 Southwark Bridge, London 
ell SHS, UK). 


© British Ornithologists’ Club 1998 


APPENDIX 


Measurements of Caprimulgus candicans at RNBM, September/December 1995 & July 
1997. 

Mean, followed in parentheses by range, standard deviation (SD) and sample size (n); 
and separated by a comma from data for the immatured plumage described in this note. 
The measurements of this bird are excluded from the mean, range and standard deviation 
calculations. All measurements are in millimetres, apart from weight in grams. 

Weight 49.3 (52-46, SD 2.66, n=6), 45; Wing chord 148 (145-150, SD 1.94, n=6), 
149; Maximum flattened wing 151 (146-154, SD 2.19, n=7), 150; Tail (from base of 
central rectrices) 104 (98-107, SD 3.69, n=6), 93, Tarsus (to extreme rear of joint) 24.9 
(24.1-—25.8, SD 0.6, n=6), 25; Culmen (from anterior of nostrils to tip of bill) 6.5 (6.3-6.6, 
SD 0.1, n=5), 5.7; Bill width (at posterior edge of nostrils) 5.7 (5.3-6.0, SD 0.25, n=5), 
5.4. 


IN BRIEF 


The eggs of the Pink-headed Duck 


by Michael Walters 
Received 7 Fuly 1997 


In his paper on the phylogeny of the pochards, Livezey (1996) uses a 
large number of morphological characters to create a new phylogeny 
which, among other things, regards the extinct Pink-headed Duck 
Rhodonessa caryophyllacea as forming a duotypic genus with the 
Red-crested Pochard Netta rufina. Among the related characters 
considered, Livezey includes nest site and clutch size, but not egg 
shape, size or colour. Shape and size are readily available from 


Schénwetter (1960). Egg colour has had a bad press since Lack’s (1958) 


In Brief 188 Bull. B.O.C. 1998 118(3) 


paper on this character in the Turdidae, but in fact the Turdidae are in 
many ways aberrant in this respect, and what is true for that family 
should not be assumed to be true for the entire class Aves. 

It is, perhaps, not well-known that the eggs of Rhodonessa 
caryophyllacea are extraordinary, being quite unlike those of any other 
member of the Anseriformes. They are stone-white in colour (though 
sometimes nest-stained), almost spherical in shape, and with a matt 
surface or a slight gloss, but lack the ‘“‘soapy’’ surface texture 
characteristic of most duck eggs. By contrast, the rest of the Aythyini 
(as defined by Livezey) have eggs of the usual duck shape, varying from 
creamy to dark olive green but never white. Because of its rather 
limited range, egg colour is more useful in indicating differences than 
relationships (unrelated birds can produce similar eggs), but given the 
general uniformity of duck eggs, the bizarre appearance of the eggs of 
Rhodonessa caryophyllacea seems not insignificant. 

For convenience, I here give a breakdown of the sizes and colours of 
eggs of the species covered by Livezey, the former from Schénwetter, 
the latter from my observations of eggs in The Natural History 
Museum at Tring (Table 1). The latter may hold the only known eggs 
of Rhodonessa caryophyllacea, including those seen and measured by 
Sch6nwetter; he states that he saw the eggs from the “British Museum’’ 
(i.e. Hume collection) and the Stuart Baker collection. Although the 
Pink-headed Duck was kept in captivity in a number of places in 
Europe, it never made any attempt to breed in captivity (Delacour 
1956, p. 199). Average measurements are in parentheses, sample sizes 
in brackets. Egg colours are given in layman’s terms rather than 
colour-coding owing to the large number of variations in shade which 
occur, but are not relevant here. 

The nest of the Pink-headed Duck was probably also unusual. F. A. 
Shillingford (in Baker 1908, p. 50-56) was the only person who 
described the nest, on which Baker commented and bemoaned that the 
finder did not give more details. The nest was concealed in the middle 
of big tufts of grass with a few feathers but, unlike most duck’s nests, 
was not lined. As doubt has been expressed (L. F. Kuff im litt) regarding 
the identity of the known eggs of the Pink-headed Duck, it may be as 
well to comment on this point. Although lacking the “‘soapy”’ surface 
texture of most duck eggs they have a faintly oily translucence and I am 
satisfied that they are anatid eggs. They are totally unlike the opaque 
eggs of owls, raptors, or any other egg that I have ever seen. 

Details of the eggs in the collection of The Natural History Museum 
are as follows (see also Knox & Walters 1994): 


1891.3.20.6019-22. 4 eggs out of a clutch of 9, collected by T. Hill on 
3 July 1880, at Purneah, India. The clutch was well incubated. These 
eges were obtained by F. A. Shillingford from Hill, and passed to the 
Hume collection. There is no reason to suppose from Hume & 
Marshall (1879-81) that any more of this clutch still exists. The eggs 
have large side-blown holes, and the others may well have broken in 
blowing. Measurements: 6019: 45.5 x 41.0. 6020: 44.95 x 42.26. 6021: 
43.35 x 42.24. 6022: 46.3 x 40.95 mm. 


Bull. B.O.C. 1998 118(3) 


189 


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1891.3.20.6018. One egg, collected by F. A. Shillingford; Hume 
collection (no date) at Zillah, Purneah, India. Measurements: 
46.0 x 44.06 mm. 

1925.12.25.5391. One egg,|\obtained by» Lt: ,\H. &. Barnesifreomtae 
Indian Museum, Calcutta, and passed to the J. Davidson collection. It 
is without locality or date, but has the name Reilly on it in 
pencil—presumably the collector. There is no reason to believe that 
this egg was in any way associated with any of the other known 
specimens. Measurements: 47.0 <x 44.9 mm. 

1925.12.25.5392. One egg, without date or locality. Davidson’s MS 
catalogue states: ““Sent to me by Mr. Irwin. It: is one of the clutch 
mentioned in Hume & Marshall’s ‘Game Birds of India’ as taken by 
Mr. Hill in either Malda or Purnesh [sic].’’ The Mr. Irwin is probably 
Valentine Irwin (1838-73) who was in the Indian Civil Service 
1862-73. Davidson would not have seen the eggs to which he refers, 
but it is certain that Irwin’s statement is incorrect, as this egg differs 
from those in being end-blown, and was evidently not from a 
well-incubated clutch. However, the statement of Davidson misled a 
previous curator, the late Shane Parker, into believing that all known 
eggs of the species were from the same clutch. Measurements: 
50.2 X 49.8 mm. 

1973.32:). One egg im the Stuart Baker collection) jeslleerea at 
Maldah, East Bengal, in June 1898, by “J. Shillingford” (?error). 
Measurements: 46.95 x 44.6 mm 


Hume (in Hume & Marshall 1879-81, p. 173-180) expressed the 
opinion that Rhodonessa was very closely related to Anas, and that he 
would have merged it but for the very extraordinary eggs. However, 
Johnsgard (1961) believed the species to be closely related to the 
pochards, while Wolfenden (1961) and Brush (1976) supported this 
view from examination of the osteology and feather protein analysis 
respectively. Its remarkable eggs would suggest that it was not closely 
related to any other duck and possibly represented a relic of an old line 
that had died out elsewhere. While it is not suggested that the 
nidification is preeminent, it is surely a factor which should at least be 
taken into consideration. 


Acknowledgements 


I am grateful to Lloyd F. Kiff for his valuable comments on an earlier draft of this paper. 


References: 

Baker, E. C. S. 1908. The Indian Ducks and their Allies. Originally published as part of 
a series of papers in 7. Bombay Nat. Hist. Soc. 11: 185-191. Reprinted as “The 
Indian Ducks and their Allies’? (1908), and rewritten almost without alteration in 
Game Birds of India, Burma and Ceylon (1921), vol. 1. 

Brush, A. H. 1976. Waterfowl feather proteins: analysis of use in taxonomic studies. 
J. Zoology 179: 467-498. 

Delacour, J. 1956. The Waterfowl of the World. Vol. 2. Country Life Ltd, London. 

Hume, A. O. & Marshall, C. H. T. 1879-81. The Game Birds of India, Burmah and 
Ceylon Vol. 3. Calcutta. 

Johnsgard, P. A. 1961. The tracheal anatomy of the Anatidae and its taxonomic 
significance. Ann. Rep. Wildfowl Trust 12: 58-69. 


In Brief 191 Bull. B.O.C. 1998 118(3) 


Knox, A. G. & Walters, M. P. 1994. Extinct and Endangered Birds in the collections of the 
Natural History Museum. Brit. Orn. Club Occ. Pub. no. 1. British Ornithologists’ 
Club, Tring 

Lack, D. 1958. The significance of colour of Turdine eggs. [bis 100: 145-166. 

Livezey, B. C. 1996. A phylogenetic analysis of modern pochards (Anatidae: Aythini). 
Auk 113: 74-93. 

Schonwetter, M. (ed. W. Meise) 1961. Handbuch der Oologie Lieferung 3: 129-131. 
Akademie-Verlag, Berlin. 

Wolfenden, G. E. 1961. Postcranial osteology of the Waterfowl. Bull. Florida State Mus. 
Biol. Sci. 6: 1-29. 


Address: M. P. Walters, Bird Group, The Natural History Museum, Akeman St., Tring, 
Herts HP23 6AP, U.K. 


© British Ornithologists’ Club 1998 


‘Dark-morph’ Sharp-shinned Hawk reported 
from California is normal juvenile female of 
race perobscurus 


by William S. Clark & Brian K. Wheeler 
Received 2 August 1997 


Patten and Wilson (1996) report an alleged dark-morph Sharp-shinned 
Hawk Accipiter striatus from California. The hawk described and 
depicted is, however, a perfect example of a darker juvenile female of 
the race perobscurus, fitting Friedmann’s (1950) description, and 
identical to three juvenile female specimens of hawks of this race from 
the American Museum of Natural History (Fig. 1). The authors state 
that their dark hawk was not an individual of this race, but they provide 
no supporting arguments. Further reasons why the hawk described and 
depicted in Patten & Wilson (1996) is not a dark morph of the 
Sharp-shinned Hawk are enumerated below. 

Neither dark-morph nor melanistic (see below for differences) 
individuals would show pale superciliaries, as shown clearly in the 
hawk in question. 

Dark-morph individuals of Accipiter (striatus) ventralis from the 
Andes are quite different from both the photograph and the description 
of this bird. They have dark grey underparts with a variable amount of 
rufous and lack pale supercilia. 

Dark-morph individuals in two African species, A. tachiro and A. 
ovampensis, are uniformly dark: black in adults and uniformly dark 
brown in juveniles, with perhaps some rufous streaking (as shown in 
del Hoyo et al. (1994) and Zimmerman et al. (1996)). They lack pale 
superciliaries. 

A dark colour morph has not been described for this species. To be 
valid, such a morph should be described in more detail and should 
include many photographs of a bird in hand or a collected specimen. 


=" 


In Brief 192 Bull. B.O.C. 1998 118(3) 


Figure 1. Three female and one male Accipiter striatus perobscurus, all juveniles. 


Dark-morph individuals usually show all dark underwing coverts and 
lack pale supercilia. 

We have no arguments with the interesting discussion of 
dichromatism in raptors that follows the brief description of the hawk 
in question. 

According to ‘Terres (1980), ‘dark morph’ is a regularly occurring 
colour morph, whereas ‘melanistic’ is an abnormal plumage. As this 
species does not have a described dark morph, the hawk referred to in 
this article is therefore an alleged melanistic. 

The specimens of A. s. perobscurus shown in Figure 1 were collected 
in the breeding range of this race, the humid temperate forests of 
coastal Washington and British Columbia, to the north of the 
California location of the hawk in question. This darker coloration, no 
doubt a perfect example of Gloger’s Rule, is shown on several other 
raptor species in this range, e.g. Merlin Falco columbarius suckleyi and 
Northern Goshawk Accipiter gentilis laingi. Dark Merlins are regularly 
recorded in California. 

It is not clear why there are no official records of A. s. perobscurus 
from California, as a juvenile of this race, found dead in southern 
California, was prepared as a specimen and deposited in the collection 
of Rio Hondo College in Whittier by John Schmitt (pers. comm.), and 
a specimen of a juvenile male of this race in the National Museum of 
Natural History was collected in Lassen Park, California, in 1908. 

In summary, it is most likely that the hawk reported in Patten & 
Wilson (1996) is a darker female of the race Accipiter striatus 


In Brief 193 Bull. B.O.C. 1998 118(3) 


perobscurus, which has occurred at least twice in California. It is most 
certainly not melanistic. 


References: 

del Hoyo, J., Elliot, A. & Sargatal, J. (eds) 1994. Handbook of the Birds of the World. 
Vol. 2. Lynx Edicons, Barcelona. 

Friedmann, H. 1950. The birds of North and Middle America, Part XI. Bull. U.S. Nat. 
Mus. 50. 

Patten, M. A. & Wilson, J. C. 1996. A dark-morph Sharp-shined Hawk in California, 
with comments on dichromatism in raptors. Bull. Brit. Orn. Cl. 116: 266-270. 
Terres, J. K. 1980. The Audubon Society Encyclopedia of North American Birds. Knopf, 

New York. 
Zimmerman, D. A., Turner, D. A. & Pearson, D. J. 1996. Birds of Kenya and Northern 
Tanzania. Princeton University Press. 


Addresses: William S. Clark, 7800 Dassett Court, Apt. 101, Annandale, VA 22003, 
U.S.A. Brian K. Wheeler, 1071 Ponderosa Circle, Longmont, CO 80501 U.S.A. 


© British Ornithologists’ Club 1998 


The identity of Kos histrio challenger 
Salvadori 1891 


by Fon Riley, Damien Hicks & Fames C. Wardill 
Received 21 August 1997 


The Red-and-blue Lory Eos histrio is endemic to the Sangihe and 
Talaud islands, north of Sulawesi, Indonesia, and according to the 
literature there are three subspecies, the nominate, talautensis and 
challengert. ‘Vhe distinction between them has been based on the extent 
of black in the wings, differences in the blue breast band, and size. E. h. 
histrio (Statius Muller, 1776) is endemic to the Sangihe islands, where 
it is historically known from Sangihe, Siau and Ruang (White & Bruce 
1986) and Tagulandang (Meyer & Wiglesworth 1898). The nominate 
subspecies has more black on the wings in comparison to talautensis; 
the secondaries are red with black terminal edging, 7 to 12 mm wide in 
adults; the greater wing coverts are tipped black, forming a bar, with a 
second black bar on the median wing coverts, and the first 3 external 
primaries are black (Meyer & Wiglesworth 1894). 

E.h. talautensis Meyer & Wiglesworth 1894 is endemic to the ‘Talaud 
islands, where it is historically known from Karakelang, Salibabu and 
Kabaruan (White & Bruce 1986). This subspecies has more red on the 
wings in comparison to the nominate; the secondaries are red with 
black terminal edging 2 to 5 mm wide in adults, the wing coverts are 
almost entirely red, and the first and third external primaries are edged 
broadly with red (Meyer & Wiglesworth 1894). 

E.. h. challengeri Salvadori 1891 was reported to be from the Nenusa 
islands, located approximately 35 km to the north-east of Karakelang, 


In Brief 194 Bull. B.O.C. 1998 118(3) 


or the Miangas islands, located approximately 100 km to the north of 
Karakelang (Meyer & Wiglesworth 1898). Meyer & Wiglesworth 
(1898) discussed this form, having examined one of the three type 
specimens. ‘The specimen was smaller and with the blue breast band 
less extensive and more or less mingled with red. 

During the course of research arising from Action Sampiri, a 
conservation project based on the Sangihe and Talaud islands (Riley 
1997), we have investigated the identity of challengeri and it is our 
conclusion that it is not a valid subspecies. It is apparently only known 
from the type specimens, although there are claims that the subspecies 
was observed in 1993 on 'Talaud (Nash 1993). The type specimens were 
obtained by the Challenger expedition on 10 February 1875: 
‘“... between the Meangis and Tulur or Talaur [Talaud] Islands, south 
of the Philippines. ‘The ship was nearest to the Island of Kakarutan, of 
the Meangis group. The large hilly island of the Talaur Group, 
Karakelang, was seen in the distance, covered with forest, but with 
numerous patches of cultivation. A canoe, sharp at both ends and 
without outriggers, of the Ke Island build, manned by 22 men and 
boys, came off to the ship... brought mats and very pretty blue 
and red Lories for sale. The birds were secured to sticks by means of 
rings made of cocoa-nut shells as at Amboina...’’ (Moseley 1879). 

Much confusion has arisen in the literature as to exactly which group 
of islands the type specimens are allegedly from. In the above quote, 
Moseley misnames the Nenusa islands as the Meangis islands, and this 
error has been perpetuated e.g. White & Bruce (1986), Forshaw (1989), 
Nash (1993), who all state challengeri is from the Miangas islands. 

‘The Nenusa islands should be the purported type locality: Kakarutan 
is an island in the Nenusa islands and Karakelang would not be visible 
from any island in the Miangas, but would be seen clearly from 
Nenusa. 

However, because the type specimens were brought to HMS 
Challenger by local fishermen from the island, and no lories were 
observed in the wild on the Nenusa group, there is no direct evidence 
that the birds were native to Kakarutan. These circumstancs therefore 
cast serious doubt on the Nenusa islands being the type locality. 

It is our opinion that the specimens described as challengeri are in 
fact immature or sub-adult talautensis. The two key characteristics on 
which the subspecies is described, its small size and plumage variations, 
can be shown not to exclude entirely either /istrio or talautensis 
subspecies. 

‘The measurements presented in White & Bruce (1986) and Forshaw 
(1989) are misleading, as immature and adult /istvio and immature 
talautensis have measurements overlapping or approaching those of 
challengeri (Meyer & Wiglesworth 1898, see Table below). ‘Therefore 
size alone cannot be used to separate challengert. 

Immature talautensis are very similar in size and colouration to 
challengeri, particularly in having the blue breast band mixed with red; 
this fact was noted by Mivart and depicted in his Plate VII (1896). 
Whether the type specimens are adult has been questioned by other 
authors (Meyer & Wiglesworth 1898). Recent observations by the 


In Brief 195 Bull. B.O.C. 1998 118(3) 
TABLE 1 


Measurements of specimens of Eos histrio showing subspecific size overlap 


Subspecies Age Number Wing length (mm) Reference 

histrio Adult 14 154-178 Meyer & Wiglesworth 1898 
histrio Adult 14 160-181 Forshaw 1989 

histrio Immature 1 154 Meyer & Wiglesworth 1898 
talautensis Adult 17 163-171 Meyer & Wiglesworth 1898 
talautensis Adult 18 160-173 Forshaw 1989 

talautensis Immature 1 LS Meyer & Wiglesworth 1898 
““challengert’ ‘‘Adult”’ 1 152 Meyer & Wiglesworth 1898 
‘challenger?’ ‘‘Adult”’ 3 152-156 Forshaw 1989 


authors, of captive birds on Talaud, suggest that immature birds vary 
considerably in the extent and intensity of blue on the breast and the 
amount of blue on the crown. Some birds have been observed with an 
ill-defined blue breast band mixed with red and only traces of blue on 
the crown. In the case of some individuals, much of the blue breast 
band is lost altogether; the reported sightings of captive challengeri on 
Talaud in 1993 are thought to relate to such birds. 

Red-and-blue Lories are extremely common cage birds on Sangihe 
and Talaud. This has probably been the case for a long time, and it 
is our opinion that the ‘“‘challengeri’’ specimens were brought to 
Kakarutan from Karakelang as cage birds, and then traded to HMS 
Challenger. It is perhaps significant that there is no record of the ship 
having docked in the Talaud islands (Moseley 1879), where further lory 
specimens would undoubtedly have been available. 

The Nenusa islands are located less than 40km offshore of 
Karakelang. We question if this narrow strait is a sufficient barrier to 
the movement of Eos histrio between the two island groups. It has been 
reported that large flocks of Eos histrio used to roost on the small Sara 
islands (Hickson 1889) and movements of lories between the Talaud 
islands are still reported by local people (Riley 1995). 

Residents of Nenusa (and the Miangas islands) interviewed by us in 
1995 and 1997, reported never having seen, or heard of E. histrio 
occurring on their islands. In all other locations within the Talaud 
islands, local residents have a sound knowledge of the birds of their 
environs and in particular the Red-and-blue Lory, which is an 
attractive and easily identified species. If the birds had ever occurred on 
Nenusa, an oral record would surely survive within the knowledge of 
native islanders. 

No ornithological fieldwork has been undertaken by us on Nenusa or 
Miangas, and surveys have not been encouraged due to the 
inaccessibility of both island groups. The islands are reported to be 
now almost entirely covered in agricultural plantations and scrub and 
there are no extensive areas of forest. It is thought likely that there is 
little or no suitable habitat to support populations of EF. histrio, but 
brief surveys are needed. 


In Brief 196 Bull. B.O.C. 1997 118(3) 


Acknowledgements 


The authors would like to thank the following organisations who provided financial 
sponsorship for Action Sampiri: British Petroleum, BirdLife International, Flora & 
Fauna International, Oriental Bird Club, The Parrot Society, Percy Sladen Memorial 
Fund, The University of York, Stiftung Avifauna Protecta, Royal Geographical Society 
and British Ornithologists’ Union. Research in Indonesia was conducted jointly with 
Universitas Sam Ratulangi, Manado, Sulawesi and under the sponsorship of Pusat 
Penelitian dan Pengembangan Biologi, Bogor and Lembaga Ilmu Pengetahuan 
Indonesia. 


References: 

Forshaw, J. M. 1989. Parrots of the World. Blandford Press, London. 

Hickson, S. J. 1889. A Naturalist in North Celebes. Murray, London. 

Meyer, A. B. & Wiglesworth, L. W. 1894. Ueber eine erste Sammlung von Vogeln von 

den Talaut Inseln. 7. Orn. 42: 237-253. 

Meyer, A. B. & Wiglesworth, L. W. 1898. The Birds of Celebes and Neighbouring Islands. 

Friedlander, Berlin. 

Mivart, St. G. 1896. A Monograph of the Lories, or brush-tongued parrots, comprising the 

family Loridae. R. H. Porter, London. 

Moseley, H. N. 1879. Notes by a naturalist on the “Challenger”. MacMillan & Co, 

London. 

Nash, S. V. 1993. Concern about trade in Red-and-blue Lories. TRAFFIC Bulletin 13: 
93-96. 

Riley, J. 1995. Preliminary Assessment of the Status and Utilization of the Red-and-blue 
Lory Eos histrio on Talaud and Sangihe. PHPA/BirdLife International-Indonesia 
Programme-Memorandum Teknis No. 10. 

Riley, J. 1997. Biological surveys and conservation priorities on the Sangihe and Talaud 
islands, Indonesia. CSB Conservation Publications, Cambridge. 

White, C. M. N. & Bruce, M. D. 1986. The Birds of Wallacea (Sulawesi, The Moluccas 
and Lesser Sunda Islands) B.O.U. Check-list no. 7, British Ornithologists’ Union. 


Addresses: Jon Riley, 1 The Croft, North Sunderland, Seahouses, Northumberland 
NE68 7XA, U.K. James C. Wardill, 27 Carlton Avenue, Hornsea, East Yorkshire 
HU181JG, U.K. Damien Hicks, Little Wyllies, Deakes Lane, Cuckfield, West 
Sussex RH17 5JB, U.K. 


© British Ornithologists’ Club 1998 


Clarification of the type locality of the 
Golden-shouldered Parrot Psephotus 
chrysopterygius 


by Stephen Garnett 
Received 3 September 1997 


The Golden-shouldered Parrot Psephotus chrysopterygius was first 
mentioned by Gould in November 1857 (Anon. 1857; see Bruce & 
McAllan 1990) on the basis of three specimens collected by J. R. Elsey, 
surgeon and naturalist with an expedition across northern Australia 
led by A. C. Gregory. Inconsistencies between the locality and date 
published on the specimen labels in the British Museum, in a letter 


In Brief 197 Bull. B.O.C. 1997 118(3) 


from Elsey to Gould in September 1857 (McDonald & Colston 1965), 
in the original description by Gould (Gould 1857, see Schodde & 
Mason 1997) and in the official account of the expedition by the 
Gregory brothers (Gregory & Gregory 1884), have led to confusion 
about the type locality. Some publications have suggested that the type 
locality was on the Norman River south of Normanton (Chisholm 
1964, Condon 1975), others that it is southeast of Normanton in the 
vicinity of Haydon (Mathews 1916-17) and one that it was at both 
(Schodde & Mason 1997). This note aims to clarify the type locality. 

Elsey’s specimen labels (1857.9.18.51/52/53), as described by 
McDonald & Colston (1965), list two collection sites and dates: No. 52. 
Labelled as female, actually an immature male: Lat. 18°S, long. 141°E: 
14th Sept. 1856. No. 53. Female: Lat. 18°S, long. 141°30’E: 15th Sept. 
1856. No. 51. The male, and type specimen of the species, lacks its 
original label. 

In his original description Gould (1857) referred to the same date 
and location but in a new combination: 


“Mr Elsey states that they were procured on the 14th September, 1856, in lat. 18°S. 


and long 141°30’E., that their crops contained some monocotyledonous seeds, ...’’. 


His description of the three specimens together implies he thought they 
were all collected on the same occasion. Curiously Mathews (1916-17), 
quoting the same notes, is more precise, describing the locality as 18°S 
and 141°33’E. 

Excerpts from Elsey’s letter (McDonald & Colston 1965) mention 15 
and 16 September, but not the 14th, and give more detail of the 
circumstances under which at least one of the parrots was collected: 


‘““In the same way I nearly lost the beautiful Euphemza. It settled in a tree close to me 
while two of our party were out after horses, and I watched it for an hour, with finger 
on the trigger until their return, expecting it to fly any moment. [It was a recognised 
signal that a shot fired at camp recalled all members of the party.] At last I got three 
specimens, popped them into my coat and set off, and it was 12 hours after, 7 p.m., 
before I had time by firelight to skin them.” 


And in another part of the letter: 


“T saw it for two days only, Sept. 15 and 16, lat 18°N [=S], long 141°E, and procured 
three specimens.” 


As background to the movements of the exploratory party during this 
period, the Gregorys’ (1884) journal describes the three days as follows: 


“14th September (Sunday)—At 5.50 proceeded down the creek on a nearly west 
course, searching the channel in its winding course for water, but without success, till 
10.0, when we found a pool of good water fifty yards long and two feet deep, at which 
we encamped. Some blacks had been camped at this pool, and their fires were still 
burning. The country on the creek is very poor, with patches of open melaleuca scrub, 
box, bloodwood, leguminous ironbark, terminalia, white gum, and a few pandanus, 
triodia, and a little very dry grass. The soil sandstone, with ironstone gravel. The 
native bee appears to be very numerous, and great numbers of trees have been cut by 
the blacks to obtain honey. 

Lat. by a Aquilae, 17 degrees. 59 mins. 26 secs. 


In Brief 198 Bull. B.O.C. 1997 118(3) 


15th September—At 8.15 a.m. resumed our journey north 10 degrees mag., over very 
level country thinly wooded with box, bloodwood, melaleuca, terminalia, grevillea, 
and cotton-trees, also a small tree which we recognised as Leichhardt’s ‘little 
bread-tree’, the fruit of which, when ripe, is mealy and acid, but made some of the 
party, who ate it, sick. Several dry watercourses trending west were crossed, and 
2.5 p.m. camped at a small waterhole in a sandy creek, fifteen yards wide. By enlarging 
the hole we obtained, though with difficulty, a sufficient supply of water for our horses. 
On the flats near the creek the grass was good, but very dry. 

Lat. by a Aquilae, 17 degrees. 46 mins. 11 sec. 

16th September—Although our horses required a day’s rest, none of our camps for 
some days had afforded a sufficient supply of water and grass for a second night; we 
therefore continued a north 20 degrees east course at 6.25 a.m.; at 7.30 a.m. came to a 
creek which we followed east an hour and a half, when it was reduced to a small gully, 
and again steered north-north-east, passing over much poor country with patches of 
melaleuca scrub, the country perfectly level; at 2.0 p.m. came to a sandy creek which 
we followed to the west till 6.5 p.m. without any water; camped in an open grassy box 
flat; I then walked down the creek and was fortunate in finding a pool of water 
half-a-mile distant, and as soon as the moon rose we drove the horses to the water and 
filled our saddle bags. Few parts of our journey have been through country so destitute 
of animal life as the level plain we have traversed since leaving the Flinders River—no 
kangaroo or even their track; emu tracks very rare, and very few birds were at the 
waterholes. Many of the sleeping frames of the blacks have been observed, and 
thousands of deep impressions of their feet in the now dry and sun-baked clay show 
that during the rainy season the extremely level nature of the country causes it to be 
extensively inundated. 

Lat. by Capella, 17 degrees. 34 mins. 5 secs.; var. compass 4 degs. 50 mins east.” 


During September Golden-shouldered Parrots feed in the early 
morning and late afternoon, spending the first hour after dawn and 
most of the middle of the day perching quietly in trees from which they 
are reluctant to move. During September elsewhere in the species’ 
range, the morning feeding session usually takes place between about 
7.00 and 8.00 a.m., after which the birds may not move from their 
daytime roost until after 4.00 p.m. (pers. obs.). From Elsey’s descrip- 
tion, the first bird he collected is unlikely to have started feeding. ‘The 
other two birds appear to have arrived at the waterhole later than the 
first and, from the presence of seed in the crop of at least one specimen, 
were probably coming in to drink after they had finished feeding. ‘The 
only day when this could have happened was 15 September. 

At that time on the expedition the party usually set out before 
6.30 a.m. On the morning of the 15th, however, it evidently took at 
least an hour for the horses to be found and brought back to camp. 
Although Elsey implies he left at 7.00 a.m., whereas the Gregorys put 
the time at 8.15 a.m., it is more likely that Elsey shot the first specimen 
at 7.00 a.m. This is consistent with a bird perching for an hour in the 
early morning before starting to feed. By the time the expedition 
actually started travelling at 8.15 a.m. Elsey had shot two more birds 
which had finished feeding. Although the party stopped travelling in 
the early afternoon, Elsey may not have had the opportunity to skin the 
birds before the evening because of his duties as expedition doctor, 
possibly tending those who became sick after eating the fruit of the 
‘little breadfruit tree’ (Parinari nonda). 

Of the alternative dates, 14 September is unlikely because on that day 
the expedition set out long before 7.00 a.m. and, while Elsey might have 
found birds at their daytime roost after they reached the waterhole, no 


In Brief. 199 Bull. B.O C1997, 148(3) 


more travelling was undertaken that day. Birds may have been labelled 
as having been collected on the 14th because they were collected at the 
campsite of the night of the 14th, not because they were actually 
procured on that day (I. A. W. McAllan pers. comm.). The 16th is also 
unlikely because the party again started too early. Furthermore the 
gibbous moon rose that evening at 8.00 p.m. (R. Allen pers. comm.). 
From the Gregorys’ description, therefore, the party would have been 
driving their horses to water only an hour after the time Elsey stated he 
began skinning out the parrots. Also, while a 6.30 a.m. starting time of 
the 16th fits better with the starting time of 7.00 a.m. suggested by 
Elsey, it is highly unlikely that a Golden-shouldered Parrot would have 
had seed in its crop so early in the day or have left the roost site for long 
enough for Elsey to have watched it for an hour. 

On this basis the type locality matches a waterhole called Dingo Hole 
at 17°59'26"S, 141°32'57”E on Clarina Creek near the northern bound- 
ary of Gum Creek Station, which corresponds almost exactly with 
the locality quoted by Mathews (1916-17). It is not known whether 
Mathews’ accuracy was a serendipitous slip or was derived from other 
information since mislaid. This site was visited in November 1992 and 
found to be a small waterhole 20 x 20 m, surrounded by vegetation that 
still fits well the description provided by the Gregorys. ‘wo months 
earlier the waterhole would have been larger and fitted even more 
closely the 50 yard length described for September 1856. At the time 
it was visited there were no other places along Clarina Creek known to 
hold water through the dry season. The location of 18°S, 141°E, which 
is near the Norman River, appears to have been a generalisation on the 
part of Elsey and it should not be inferred that the birds were collected 
on the Norman River south of Normanton. 

The type locality of the Golden-shouldered Parrot is of particular 
interest for three reasons. First, the nearest place where the species has 
been recorded since 1856 is over 200km away. Secondly, Elsey’s 
remark that he saw the birds over two days suggests that the three birds 
he shot were not an isolated group but part of a larger population, now, 
apparently, extirpated. Thirdly, the suggestion that the birds were 
collected further west has led to potential errors in biogeographic 
analysis of the distribution of tropical granivores in Australia (D. 
Franklin pers. comm.). 


Acknowledgements 


I am grateful to John Woodburn for his hospitality during our stay in Normanton in 
1992, to the owners of the Gum Creek and Hereford pastoral leases for allowing me entry 
to their properties, to Norman Wettenhall for access to his peerless library, to Ian 
McAllan for commenting on the manuscript, to Les Christidis for help with determining 
the time of moonrise on Cape York Peninsula 142 years ago and to Hugo Phillipps for 
retrieving final references. 


References: 

Anon. 1857. Zoological Society. The Literary Gazette 2130: 110. 

Bruce, M. D. & McAllan, I. A. W. 1990. Some problems in vertebrate nomenclature. II 
Birds, Part 1. Boll. Mus. reg. Sci. nat. Torino 8: 453-485. 


Books Received 200 Bull. B.O.C. 1998 118(3) 


Chisholm, A. H. 1964. J. R. Elsey, surgeon, naturalist, explorer. Queensland Naturalist 
17: 60-70. 

Condon, H. T. 1975. Checklist of the Birds of Australia. Part 1. Non-Passerines. RAOU, 
Melbourne. 

Gould, J. 1857. On several new species of birds from various parts of the world. Proc. 
Zool. Soc. London 25: 220-222. 

Gregory, A. C. & Gregory, F. T. 1884. Journals of Australian Explorations. James & Beal, 
Brisbane. 

McDonald, J. D. & Colston, P. R. 1965. J. R. Elsey and his bird observations on 
Gregory’s overland expedition, Australia, 1856. Emu 65: 255-278. 

Mathews, G. M. 1916-17. The Birds of Australia. Vol. 6. Witherby & Co., London. 

Schodde, R. & Mason, I. J. 1997. Aves (Columbidae to Coraciidae). In W. W. K. 
Houston & A. Wells (eds), Zoological Catalogue of Australia. Vol. 37.2. CSIRO 
Publishing, Melbourne. 


Address: Stephen Garnett, Department of Environment, P.O. Box 2066, Cairns, 
Queensland 4870, Australia. 


© British Ornithologists’ Club 1998 


BOOK RECEIVED 


Gatter, Wulf 1997. Birds of Liberia. Pp. 320, 36 plates in colour, 8 monochrome plates, 
maps and text figures. Pica Press. ISBN 1-873403-63-1. £40. 225 x 290 mm. 


This is the fifth book to provide a detailed checklist of the birds of a West African 
country, the previous four all being in the BOU Checklist series. It is a beautifully 
produced book such as we have come to expect from Pica Press and is exceptionally well 
illustrated with painted portraits of 29 species, and coloured photographs of 56 species. 
In addition there is an invaluable collection of 30 coloured photographs of habitats, most 
half page, and 21 assorted monochrome photographs. 

The checklist comprises 77 pages of introductory text, a systematic list (pp. 91-246) of 
615 accepted species and bracketed entries for a further 14 species of doubtful validity, 
followed by six appendices in 20 pages, a two page gazetteer, four pages of references and 
an index of scientific and English names. Gatter’s checklist covers the history of 
ornithology in Liberia, geology and topography, climate, vegetation zones and habitats of 
importance to birds. Then follow detailed essays of breeding seasons, moult, migration 
and ecology of forest birds, including mixed species flocks. The introduction finishes with 
short accounts of biogeography and conservation. Liberia is the only West African 
country located almost entirely within the forest zone. Gatter, a professional forester, 
gives a very clear account of the forest vegetation and makes a major contribution to our 
knowledge of the ecology of forest birds, with a wealth of information on foraging heights 
and membership of mixes species flocks. However, the chapter on migration is not so 
convincing, with too much made of limited data in some cases. 

Each checklist entry has the scientific and English name of the bird mainly from Birds 
of Africa, abbreviated status as in the BOU checklists, and the number used in the two 
volumes of Mackworth-Praed & Grant (1970 and 1973). Then follows a general 
statement on abundance, status, habitat and distribution within Liberia, and migration 
relative to Liberia. Habits, ecology and annual cycle are then treated in more detail where 
data allow and for most resident breeders and suspected breeders there is a distribution 
map. The accounts of forest species in particular are good, often providing numerical 
details to back statements on seasonality etc. Most of these data come from Gatter’s own 
studies at Zwedru during 1981-84. 

References seem particularly thorough for German literature but perhaps less so for 
English publications. Some references cited in the text are missing from the bibliography 
(e.g. Bremer 1989, Pearson & Backhurst 1986), whilst others in the bibliography are not 
cited in the text (e.g. Walsh & Walsh 1976). The index of scientific and English names 
covers only the checklist entries. 

This is an important contribution to the ornithology of West Africa which will also be 


of considerable interest to all students of tropical forest birds. 
J. Frank Walsh 


NOTICE TO CONTRIBUTORS 

Bikers a are invited from Club Members or non-members, especially on taxonomic 
and distributional topics; descriptions of new species are especially welcome and 
may be accompanied by colour photographs. Two copies of manuscripts, typed on 
one side of the paper, double spaced and with wide margins, should be sent to 
the Editor, Prof. Chris Feare, 2 North View Cottages, Grayswood Common, 
Haslemere, Surrey GU27 2DN, UK. All contributions, including In Brief 
articles, should follow the style of main papers in this issue of the Bulletin. 


A contributor is entitled to 10 free offprints (16 if 2 or more authors) of the pages of the 
Bulletin in which his/her contribution, if one page or more in length, appears. Additional 
offprints or offprints of contributions of less than one page may be ordered when the 
manuscript is submitted and will be charged for. Authors may be charged for proof corrections 
for which they are responsible. 


MEMBERSHIP 

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and to receive (postage free) four quarterly issues of the Bulletin, and the annual 
index, for an annual subscription of £12 (or U.S. $26). Applications, enclosing the 
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NON-MEMBER SUBSCRIBERS & APPLICATIONS FOR BACK 
NUMBERS OR OTHER PUBLICATIONS 

The Bulletin (for 1998 onwards), together with annual index, may be purchased 
(postage free) by Non-member Subscibers on payment of an annual subscription of 
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and runs of back numbers of the Bulletin, and also other BOC Publications may 
similarly be obtained, on request to him. 


PAYMENTS 

All amounts quoted are net and should be paid in £ sterling, if possible. Payments 
in other currencies must include a further £4 for UK bank charges (except for 
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made payable to the British Ornithologists’ Club. If preferred, remittances may be 
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CORRESPONDENCE 
Correspondence on membership, changes of address and all other matters should 
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inside front cover. 


Registered Charity No. 279583 


CONTENTS 


Page 
CLUB IN@EICES. 20. ep ae ie It 129 
RASMUSSEN, P. Cc. A new Scops-owl from Great Nicobar Island ......... 141 


HORNE, J. F. M. & SHORT, L. L. A case of ‘song-capture’: Rufous-naped Lark 

uses nightjar song in place of its own song....... .... .. 2. ae eee 153 
BEGAZO, A. J. & VALQUI, T. H. Birds of Pacaya-Samiria National Reserve with 

a new population (Myrmotherula longicauda) and new record for Peru 


(Hylophilus semicinereus) . ..00. 6 6 2 oj se oe oe ss os 159 
ARCHER, A. L. & ILES, D. B. New bird records for Unguja (Zanzibar Island) 166 
COLLINS, C. T. & ARAYA, T. A. Natal pterylosis of tanagers II: Tachyphonus, 

Ramphocelus and Tangara .. 2.2... i f2. ss. .>. « ghee 12 
ANDERSON, D. L., BONTA, M. & THORN, P. New and noteworthy bird records 

from Honduras 0.5. 6.6 cu os a i So ae es ees a 178 
LANUS, B. L., CLAY, R. P. & LOWEN, J.C. A new plumage of the White-winged 

Nightjar Caprimulgus candicans (Aves: Caprimulgidae)............. 183 
In Brief WALTERS, M. ‘The eggs of the Pink-headed Duck............. 187 


CLARK, W. S. & WHEELER, B. K. ‘Dark-morph’ Sharp-shinned 
Hawk reported from California is normal juvenile female of 


race Perobscurus . os sh de Oo sd ee os ee 191 

RILEY, J., HICKS, D. & WARDILL, J. Cc. The identies of Eos histrio 
challengert Salvadori 1891 ...........5. = lor 193 

GARNETT, S. Clarification of the type locality of the Golden- 
shouldered Parrot Psephotus chrysopterygius.............. 196 
BOOK RECEIVED.) . <6 5 Sin 2 8 6 6 3 ow we ees 3 oe Oe 2 200 


¥ 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 
mail, after their current subscription has been paid. 


COMMITTEE 


Revd T. W. Gladwin (Chairman) (1997) Dr 
Mrs A. M. Moore (Vice-Chairman) (1997) N. 
Cdr M. B. Casement, OBE, RN TS: 

(Hon. Secretary) (1996) N. 
D. J. Montier (Hon. Treasurer) (1997) M 


Hon. Editor: Prof C. J. Feare 
Chairman of Publications Sub-committee: Dr R. P. Prys- Jones 
Publications Officer: S. J. Farnsworth 


R. P. Prys-Jones (1995) 
H. F. Stone (1995) 
Griffin (1997) 

J. Redman (1997) 

rR. E. Scott (1998) 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


a 
ut 
% 802 


2} ra ISSN_0007—1595 


Edited by 
Prof CHRIS FEARE 


Volume 118 No. 4 December 1998 


FORTHCOMING MEETINGS 


Advance notice of meeting dates for 1999. Eight meetings have been arranged for the 


following Tuesdays: 19 January, 16 March and 20 April—for details see below, 4 May 
(AGM and social evening—with informal “‘mini-talks’’), 6 July, 7 September (Alex Randall 
MP on “Bird conservation matters, as viewed from Westminster’’), 12 October and 30 


November (Richard ffrench on “Dickcissels in Trinidad”’). Details of speakers on 6 July and ~ 


12 October will be published when finalised. 


Tuesday 19 January 1999. Keith Betton will speak on “Birding by ear—a look at the 
world of strange, and not so strange bird sounds’. Keith was born and educated in 
London, where he quickly established a life-long interest in ornithology. At the early age of 12, 


he was appointed a member of the Department of Environment Committee on Bird Sanctuaries 


in Royal Parks, and was Official Observer for Bushy Park (1973-79). He has served on 
numerous ornithological committees, including Chairman Ornithology Section of London 
NHS (1984-92), President London NHS (1982-84), BTO Regional Representative Greater 
London (1981-93), and Council BTO (1987-91), also Committee BOC (1985-88). He is a 


prolific contributor to and editor of several journals. His high profile post as Head of Corporate 
Affairs, Association of British Travel Agents, involves travelling worldwide on business, but 


this now leaves him less time for these ornithological activities. His special interest is in Africa, — 


and in sound recordings, of which he has accumulated a large library of his own. 
Applications to the Hon. Secretary by 5 fanuary, please. 


Tuesday 16 March 1999. Nathalie Seddon will speak on “Birds of Madagascar”. 


Nathalie is currently a student at the Zoological Department of Cambridge University, and © 


studying for her PhD on the behavioural ecology of the Subdesert Mesite Monias benschi, in 
Madagascar. She is a veteran of several international ornithological expeditions, including one 
to western Ecuador. 

Applications to the Hon. Secretary by 2 March, please. 


Tuesday 20 April 1999. Roger Safford will speak on “Birds of Mauritius”. After gaining © 
his degree in Natural Sciences in Cambridge in 1988, Roger obtained his PhD on Conservation © 
of the forest-living birds of Mauritius, at the University of Kent, in 1994. His special interest — 


is in natural history and conservation in the western Indian Ocean and East Africa, and he has 


spent five years in the area, especially in Mauritius, and also in Madagascar. He is currently 
Tropics projects co-ordinator, Royal Holloway Institute for Environmental Research, 
University of London. As a Member African Waterfowl Census, he was National Co-ordinator, — 
for Mauritius 1990-93, and has been a Member IUCN Species Survival Commission, — 


since 1991. 
Applications to the Hon. Secretary by 6 April, please. 


Meetings are held in the Sherfield Building of Imperial College, South Kensington, London ~ 


SW7. The nearest Tube station is at South Kensington, and car parking facilities are available; a 
map of the area will be sent to members, on request. The cash bar is open from 6.15 p.m., and 
a buffet supper, of two courses followed by coffee, is served at 7.00 p.m. (A vegetarian menu 
can be arranged if ordered at the time of booking.) Informal talks are given on completion, at 
about 8.00 p.m. 


Overseas Members visiting Britain are particularly welcome at meetings. For 


——— 


details in advance, please contact the Hon. Secretary, Cdr M. B. Casement, OBE, RN, — 


Dene Cottage, West Harting, Petersfield, Hants GU31 5PA. 
Tel/Fax: 01730-825280 for late bookings and cancellations. 


© British Ornithologists’ Club 1998 


Apart from single copies made for the purposes of research or private study, or criticism or 
review, as permitted under UK law, no part of this publication may be reproduced, stored 
or transmitted in any form or by any means, except with prior permission in writing of the 
publishers, or in accordance with the terms of licences issued by the Copyright Licensing 
Agency. 


Enquiries concerning reproduction outside these terms should be sent to the Editor; for 
address see inside back cover. 


201 Bull. B.O.C. 1998 118(4) 


Bulletin of the 
BRITISH ORNITHOLOGISTS’ CLUB 


Vol. 118 No. 4 Published December 11 


CLUB NOTES 


The eight hundred and seventy eighth meeting of the Club was held on Tuesday, 15 
September 1998 at 6.15 p.m. 28 Members and 13 Guests attended. 

Members present were: The Revd T. W. GLaDwIn (Chairman), Miss H. BAKER, P. J. 
BELMAN, P. J. BULL, D. R. CALDER, Cdr M. B. CASEMENT RN, Professor R. J. CHANDLER, 
Professor R. A. CHEKE, Dr N. J. COLLAR (Speaker), J. FARNSWORTH, D. J. FISHER, F. M. 
GAUNTLETT, D. GRIFFIN, T. M. GuLiickx, C. R. HELm, T. P. INskipp, J. A. JoBLING, Dr 
C. F. Mann, D. J. Montier, Mrs A. M. Moors, R. G. Morcan, Mrs M. N. MULLER, 
Dr R. P. Prys-Jonss, N. J. Repman, Dr R. C. SELF, P. J. SELLAR, N. H. F. STONE, and 
M. P. WALTERS. 

Guests attending were: A. V. ANDREEV, Mrs J. C. BuLL, Mrs J. B. CALDER, Mrs M. H. 
GaUNTLETT, Mrs J. M. GLapwin, Ms C. Horr, Mrs C. INskipp, Mrs M. MOontier, 
P. J. Moore, M. J. SEpGEMOoRE, C. J. SEDGEMORE, Ms A. J. STATTERSFIELD and 
F. STEINHEIMER. 

On completion, Nigel Collar gave a presentation, illustrated with slides and museum 
specimens, entitled “‘Two Indian Enigmas’. 

Indian ornithological enigmas include Pink-headed Duck Rhodonessa caryophyllacea 
(probably extinct), Himalayan Quail Ophrysia superciliosa (missing since 1876), Jerdon’s 
Courser Rhinoptilus bitorquatus (rediscovered in 1986), Intermediate or Rothschild’s 
Parakeet Psittacula intermedia and Forest Owlet Athene blewittz. Recent work led by Dr 
Pamela C. Rasmussen of the Smithsonian Institution has shed light on the last two 
(Rasmussen & Collar 1988, Identification, distribution and status of the Forest Owlet 
Heteroglaux (Athene) blewitti. Forktail 14: 41-49; in press a, A major specimen fraud in 
the Forest Owlet Athene (auct.) blewitti: Ibis, in press b, Rothschild’s Parakeet Psittacula 
intermedia is a hybrid. Bull. Nat. Hist. Mus.). 

Seven dataless specimens sent from India around 1895 (the type) and 1907 (six others) 
were so clearly midway between Plum-headed Parakeet Psittacula cyanocephala and 
Slaty-headed Parakeet P. himalayana that Rothschild gave them the name P. intermedia, 
yet he, his curator Hartert and subsequent authorities resisted the notion that they were 
of hybrid origin. However, scrutiny of this and other material, and comparisons 
including Blossom-headed P. roseata and Grey-headed P. finschii Parakeets, disprove all 
six defences of specific status for intermedia, namely: (1) the specimens show uniform 
characters (they do not—one is even an immature himalayana); (2) they have a single 
origin (stuffing style and material suggest otherwise); (3) they have a non-captive origin 
(plumage abrasion and soiling suggest otherwise); (4) a 1932 description of hybrid 
cyanocephala X himalayana does not match intermedia (the account is muddled and only 
treats juveniles); (5) reported captive intermedia currently exist in Austria and India (this 
proves nothing); and (6) biochemical analysis showed the distinctness of Indian captives 
(these are cyanocephala X krameri, and anyway were inadmissibly analysed). Measure- 
ments confirm intermedia’s consistent intermediacy between cyanocephala and himalayana, 
a condition unknown in any true species. 

Mr Melville Sedgemore (present) recently crossed a male cyanocephala with a female 
himalayana expressly to determine if offspring show characters of intermedia. They do; 
and their measurements also closely match intermedia. There is thus no question that 
intermedia is a hybrid, and it is inconceivable that the specimens were not bred in 
captivity. 

The Forest Owlet is known from central India from seven old specimens, the most 
recent of which was purportedly collected by R. Meinertzhagen in Gujarat on 9 October 
1914. However, Meinertzhagen’s diary and specimen catalogue indicate that his presence 
in Gujarat on that date is implausible. Scrutiny of the specimen establishes that it was 
extensively remade, and that it has features (wings tied outside the body, cotton 
wing-stuffing, a once stretched-out neck) indicating that the true collector was J. 


Club Notes 202 Bull. B.O.C. 1998 118(4) 


Davidson. Examination of the British Museum register reveals a missing Davidson 
specimen from Maharashtra in 1884. 

The Forest Owlet has always been considered virtually identical to Spotted Owlet 
Athene brama, but evaluation shows that it strongly differs in having a faintly spotted 
crown and back, pale auriculars with no white rear border, a broad complete dark frontal 
collar, breast almost solid brown, boldly barred flanks, an unmarked white lower breast 
and belly, and prominently banded wings and tail (structurally it may merit Hume’s 
monotypic genus Heteroglaux). ‘These features, combined with greater clarity over 
records following the elimination of Gujarat, led to the bird’s rediscovery in 1997 by Dr 
Rasmussen with B. F. King and D. F. Abbot. It is, however, in serious danger from 
deforestation. 


The eight hundred and seventy ninth meeting of the Club was held on Tuesday, 13 
October 1998 at 6.15 p.m. 16 Members and 10 Guests attended. 

Members present were: The Revd T. W. GLADWIN (Chairman), Miss H. Baker, P. J. 
BeLman, P. J. BuLL, Cdr M. B. CasEMENT RN, Professor R. J. CHANDLER, Professor 
R. A. CHEKE, J. A. JoBLinc, R. H. Kerrie, Dr C. F. Mann, D. J. Montisr, R. G. 
Morean, Dr R. P. Pr¥s-Jones, R. E. Scott, Dr R. C. SELF and N. H. F. STONE. 

Guests attending were: Mrs J. C. BuLL, Mrs C. R. CasEMENT, J. H. W. Cuitty, Mrs 
D. Cuitty, Mrs J. M. Giapwin, Dr B. HATCHWELL (Speaker), Ms C. Horr, Mrs 
M. Monttsr, J. SCHARLEMANN and F. STEINHEIMER. 

Dr Ben Hatchwell subsequently gave a highly interesting presentation, illustrated with 
slides, entitled “‘“The Cooperative behaviour of Long-tailed Tits’. 

The Long-tailed Tit Aegithalos caudatus is one of about 300 bird species known to 
breed cooperatively—some individuals forego personal reproduction and instead help to 
rear offspring which are not their own. Such behaviour poses a problem for evolutionary 
biologists because animals are expected to act in their own selfish interests rather than in 
the interests of others. What are the benefits of cooperation and why has it evolved in 
Long-tailed Tits? 

Long-tailed Tits spend most of the year (June—February) in flocks of 10-15 birds, 
typically comprising a family of 8-10 juveniles, their parents and a variable number of 
helpers. Flocks split up in early spring, males adopting part of the flock range, while 
females either pair up‘within the flock, or disperse to neighbouring flock ranges. All birds 
start the season breeding in pairs, but 80% of nests fail due to predation, mostly by 
corvids. Pairs who fail have two options: (i) they may re-nest if failure occurs early in the 
season, or (ii) if they fail after early May, they may become helpers at another pair’s nest, 
assisting them by feeding their nestlings. About 50% of nests have at least one helper 
(maximum 4) for part of the nestling period. DNA fingerprinting and genealogical data 
indicate that helpers assist close relatives. 

Helpers benefit in two ways from their cooperation. First, by assisting relatives they 
gain indirect fitness benefits by increasing the production of related offspring. The 
presence of two or more helpers increases the amount of food given to nestlings, so each 
fledgling is heavier and has a higher probability of surviving to breed. Secondly, by 
helping, failed breeders might increase their own chance of reproducing in the following 
season (a direct fitness benefit) because helpers tend to have a better chance of surviving 
to the next season (65%) than do failed breeders who do not help (45%). 

Given these potential benefits, why has cooperative behaviour evolved so rarely? Do 
cooperative species differ ecologically from non-cooperative species? The Ecological 
Constraints Hypothesis attributes cooperation to constraints (e.g. a lack of vacant 
territories or breeding partners) which restrict dispersal, forcing grown offspring to ‘stay 
at home’ and help their parents, even though they would prefer to breed independently. 
Long-tailed Tits do not fit easily into this hypothesis because analysis of BTO ringing 
data shows that their natal dispersal does not differ from ecologically similar 
non-cooperative species. Furthermore, all birds are able to breed independently. 
Nevertheless, one can speculate on those aspects of Long-tailed Tit biology that might 
promote cooperation. First, dispersal occurs unusually late (eight months after fledging), 
giving family members ample opportunity to learn who their relatives are, enabling them 
preferentially to assist kin following breeding failure. Second, late dispersal may result 
from their small size which makes flock membership critical for winter survival. Failed 
breeders who cooperate might ‘buy’ access to the communal roosts which are 
characteristic of this species. 


Club Notes 203 Bull. B.O.C. 1998 118(4) 


Such species-specific explanations are unsatisfactory because they obviously do not 
apply to all cooperative breeders. However, they do serve to identify factors (e.g. the 
benefits of group living) which are likely to be of general significance in explaining this 
unusual behaviour. 


The B.O.C. Archive 


The British Ornithologists’ Club has been in existence now for over a hundred years, 
during which time it has not had a permanent home. A few years ago a tin box was 
returned to the Secretary from the London Zoological Society where it had been stored. 
This contained the minute books and attendance records from the earliest time (less one 
attendance “‘lost on the Underground’’!). 

It was thought an ideal time to think of the future of this material plus that of the 
correspondence etc. that had accrued over the years from various Chairmen and 
Secretaries. Mr Ronald Peal and Mrs Amberley Moore were instrumental in passing 
material to me as well as giving good advice, as did Dr James Monk. 

The archive consists, in addition, of the Herbert Stevens Archive, historical material 
brought together when a history of the Club was considered (this includes an audio tape), 
a bound set of the Bulletin and copies of the Club’s publications. A recent addition, 
provided by Mr David Griffin, is a list of speakers and subjects compiled from the 
Bulletin. 

The committee decided that the material available should be catalogued and organised 
as an official archive and I was asked to do this. The Secretaries’ and Chairmen’s 
correspondence is very limited in content; some people in the past kept every scrap of 
paper, others threw them out after dealing with them. The material that was kept does 
show the amount of work that these two posts involved, although the ability to make 
mountains out of molehills was very obvious. However it is a pity that material by such 
stalwarts as Lord Rothschild, R. Bowdler-Sharpe and W. L. Sclater were not kept. It is 
also possible that we might have had some gems by the Rev. Jourdain and N. F. 
Ticehurst. 

Some aspects of the Archive are of course ongoing, and this has meant that the index 
is in several parts to accommodate new material where necessary. The archive is at 
present housed at the Natural History Museum (Tring), thanks to the generosity of the 
museum and staff. Access is restricted, and applications to consult the archive need to be 
approved by the Chairman and Archivist; the latter will usually be able to advise on the 
material available. 

N. H. F. STONE 
Archivist 


WHY MUSEUMS MATTER: AVIAN ARCHIVES IN AN AGE OF 
EXTINCTION 


A conference organised jointly by the British Ornithologists’ Club, British Ornithol- 
ogists’ Union, The Natural History Museum and BirdLife International, 12-14 
November 1999 (see announcements in Bull. Brit. Orn. Club 118(3): 135-136 and Ibis 
140(4): 723). 


CALL FOR OFFERED PAPERS/POSTERS 


Papers/posters relevant to the conference theme are invited. Offers, with brief abstracts, 
to Dr R. Prys-Jones, Natural History Museum, Tring, Herts HP23 6AP, U.K.; fax: +44 
(0) 1442 890693; e-mail: R.Prys-Jones@nhm.ac.uk 


F.R. Lambert & P. C. Rasmussen 204 Bull. B.O.C. 1998 118(4) 


A new Scops Owl from Sangihe Island, 
Indonesia 


by Frank R. Lambert & Pamela C. Rasmussen 
Received 14 September 1998 


Situated between the Minahasa peninsula of Sulawesi and the 
Philippine island of Mindanao, the Sangihe and alaud islands (Fig. 1) 
have been poorly explored ornithologically, but are now recognised as a 
centre of avian endemism (Stattersfield et al. 1998). Recent work in the 
‘Talaud islands has revealed the presence of two new species of rail 
(Lambert 1998a, 1998b), and on Sangihe the existence of an endemic 
species of shrike-thrush Colluricincla sanghirensis has been confirmed 
(Riley 1997a, Rozendaal & Lambert in press). 

Four specimens of a scops owl collected on Sangihe Island (=Sangi 
or Great Sanghir, hereinafter “Sangihe’’) between 1866 and 1887 were 
thought not to differ from the Sulawesi Scops Owl Otus manadensis 
(Meyer 1884, Blasius 1888, Finsch 1898), and thus were never given a 
name. Meanwhile, from nearby Siau (Fig. 1), a single specimen 
obtained in 1866 was described as Scops siaoensis on the basis of its 
exceptionally small size (Schlegel 1873, Plate 4). That name was later 
synonymised with manadensis (Meyer 1884, Meyer & Wiglesworth 
1898), a treatment followed uncritically by most subsequent authors. In 
Marshall’s (1978) treatise on Asian Otus, no firm decision was taken on 
the systematic position of either szaoensis or the Sangihe population due 
to the lack of data on vocalisations. Marshall (1978) mentioned 
unspecified differences in colouration and tarsal feathering between 
manadensis and populations from islands off Sulawesi, including 
Sangihe, but he apparently examined no specimens from the Sangihe 
group. This led to his very tentative allocation of all these populations 
to the widely distributed, highly variable Moluccan Scops Owl Otus 
magicus (sensu Marshall 1978, Marshall & King 1988). Bruce (7m White 
& Bruce 1986) thought specimens from Sangihe showed affinities with 
manadensis, but the population was nevertheless allocated (op. cit.) with 
a query to magicus. In 1985, F. G. Rozendaal heard an Otus calling on 
Sangihe and collected a single specimen, noting that its call was “‘the 
same whistle as heard on ‘mainland’ Sulawesi” (F. G. Rozendaal, in 
litt. 1998). On this basis, the Sangihe bird was placed in manadensis by 
Marshall & King (1988), who reasoned (apparently without reference to 
specimens or recordings) that all taxa from islands closer to Sulawesi 
than is Sangihe (including szaoensis, mendeni of Peleng I., and kalidupae 
of Tukangbesi Is.) should also belong with manadensis. 

In 1996, FRL photographed a scops owl on Sangihe (Plate 5), 
tape-recorded its vocalisations (Fig. 2), and realised that the songs 
sounded different from Sulawesi birds. Subsequently, Riley (1997b; zn 
litt. 1998) photographed and took a few measurements of a caged scops 
owl on Sangihe in poor condition and with clipped wings, but although 
the bird soon died it was not preserved as a specimen. Both Riley 


Plate 4. Sangihe Scops Owl Otus collari (lower), in comparison with a Sulawesi Scops 
Owl Otus manadensis (AMNH 298939, upper left) and the unique specimen of Otus 
[magicus] siaoensis (upper right). Original watercolour painting by J. C. Anderton. Iris 
colour of siaoensis is assumed to be yellow based on colour of most related taxa. 


Plate 5. Photographs of a live Sangihe Scops Owl on the lower slopes of Mt. Awu near 
Telawid Atas, 30 July 1996 (FRL). 


F.R. Lambert & P. C. Rasmussen 205 Bull. B.O.C. 1998 118(4) 


Figure 1. Map showing the location of Sangihe in relation to the Talaud Islands, Siau 
and the Minahasa peninsula of northern Sulawesi. 


(1997a) and B. F. King also heard and tape-recorded scops owls on 
Sangihe. While scops owls heard by Riley (1997b) were said to sound 
“Sdentical to birds heard and seen near Manado on mainland 
Sulawesi’, sonagrams from Riley’s tape show that they are 
indistinguishable from those taped by FRL. Additionally, the plumage 
characters reported by Riley (1997b) that were said to confirm the 
placement of the Sangihe bird in manadensis can equally be applied to 
some taxa of magicus. In 1998, PCR examined four Sangihe specimens, 
photographs by FRL and J. Wardill of the two above-mentioned living 
individuals from Sangihe, and specimens of all related taxa. Although 
the Sangihe bird is indeed very like some manadensis, several 
morphological differences exhibited by all specimens and photographs 
examined, as well as numerous striking vocal differences consistent 
among recordings, indicate that it is distinct at the specific level. We 
propose for it the name: 


Bull. B.O.C. 1998 118(4) 


206 


F. R. Lambert & P. C. Rasmussen 


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F. R. Lambert & P. C. Rasmussen 207 Bull. B.O.C. 1998 118(4) 
Sangihe Scops Owl Otus collari, sp. nov. 


Holotype. Staatliches Naturhistorisches Museum, Braunschweig 
(SNMB), female, from “Sangir’’ (exact locality not given, but the 
island is centred at 3°35’N, 125°32’E), collected 2 January 1887 by Dr 
C. Platen, no. 6968; specimen ‘“‘a’’ of Blasius (1888). 


Diagnosis. A drab brownish, rather small, yellow-eyed scops owl with 
a horn-brown bill, long narrow wings, a rather long tail, and small pale 
feet and claws, the latter tipped darker (Plate 4). The eartufts are 
medium-length with buff spots, black streaks, and elliptical tips; the 
pale supercilium is rather short; the face shows little contrast, the 
darkest part being between eye and bill; the upperparts have dark shaft 
streaks and are prominently spotted buff; the scapular spots are pale 
buff on the outer web with triangular black tips; the underparts have 
prominent but mostly fine black shaft-streaking and a finely 
vermiculated base pattern; the flight feathers are banded dark brown 
and buff, but the tertials are not prominently banded; the tail has 
narrow irregular dark buff bands and wider dark brown bands; and the 
tarsal feathering ends just above the tarsal joint in front and meets 
around the rear. 


Otus collari is close to certain Otus manadensis (Plate 4, n=100+), 
especially to immatures, but relative to manadensis even from a single 
locality, the degree of variability among the six individuals examined of 
collari is much reduced. On the head, the facial disk of collari is paler 
and contrasts less with rest of face, there is a definite dark-barred area 
on the rictal bristles between eye and bill, the whitish supercilium is 
somewhat less obvious and ends above the bill, and the streaks on the 
crown are usually narrower and contrast more strongly with the paler 
brown base colour. On the upperparts, collari is slightly more coarsely 
vermiculated, with generally shorter, more diffuse, less obvious 
blackish shaft streaks, the upper wing coverts are plainer and less 
pale-spotted, and the scapulars have pale buffy outer webs with 
relatively larger triangular black tips. The underparts have a finer, 
sketchier background pattern with less definite, reduced cross-barring, 
most of it stippled rather than solid, the band interstices are drab, 
stippled brown, and have reduced contrast, while most of the streaks on 
the underparts are longer and narrower, and the pale spots on the 
underparts are usually smaller, bufher, more regular, and _ less 
conspicuous. The wings of Otus collari are longer than for manadensis, 
but they are considerably narrower due to the inner primaries being 
shorter and emarginations of the outer primaries being more proximal, 
even when corrected for size (log data divided by log wing length; 
‘Table 1), and its primaries show reduced contrast due to the pale bands 
being buffy and the dark bands being browner and less blackish. The 
tail of collari is slightly longer (Table 1) and the toes and claws 
relatively weaker than in manadensis. There is very limited overlap in 
each of these plumage and mensural characters with the extremely 
variable manadensis, (in which no more than a few of these features 
occur together in the same individual), and in combination they give 
collari a more uniform, drabber appearance than for adult manadensis. 


F.R. Lambert & P. C. Rasmussen 208 Bull. B.O.C. 1998 118(4) 


See under ‘“‘Remarks”’ for additional morphological distinctions from 
other related taxa. 

Otus collart is vocally readily diagnosed by its clear, mainly 
downslurred, fluty, liquid, relatively high-pitched, long whistle; see 
under ‘“‘Voice’’ for details. 

Distribution. ‘The new species is apparently widespread on Sangihe. 
In southern Sangihe, collari has been reported from the upper slopes of 
Mt. Sahendaruman (3°31'N, 125°32'E; F. G. Rozendaal, in litt. 1998), 
the southwest coast at Manganitu (3°35’N, 125°31°E, RMNH 84653) 
and Mt. Sahengbalira (3°31’N, 125°31’E, Riley 1997b); on the 
west-central coast, from near Tahuna (3°37'N, 125°29’E; Riley 1997b); 
and in northern Sangihe, from the northeast coast at Tabukan 
(=Tabukanlama, 3°41’N, 125°33’E, Meyer 1884), and from areas 
around Telawid, on the lower slopes of Mt. Awu (3°43'N, 125°24’E, 
Riley 1997a, FRL). 

Description of the holotype. Colour comparisons were made under 
fluorescent light. Capitalised colour names and the first numbers in 
parentheses are from Smithe (1975); Munsell (1977) colour matches are 
also presented, in which the first number and letters represent the hue 
(but are interpretable only with reference to Munsell charts), the next 
increasing value or lightness, and the last increasing chroma or 
saturation. ‘The holotype was directly compared with 11 manadensis 
assembled at the National Museum of Natural History, USNM, the 
two Leiden specimens with 25 manadensis at NNM, and the Dresden 
specimen with 19 manadensis at MTD; all four Sangihe specimens were 
compared with photographs of specimens of all taxa. For the holotype, 
colorimetric readings (using a Minolta CR-221 Chromameter) were 
taken of certain relatively solid colour plumage areas larger than 3 mm 
in diameter. Three sequential measurements were taken and summary 
statistics instrumentally calculated, then summary statistics were 
calculated for three independent sets of the above, with the colorimeter 
repositioned between each triad of measurements. Values are given in 
terms of the opponent colour coordinate system (L, a, b; Graves 1997), 
in which higher values of L show increasing lightness (0-100), higher 
positive a values increasing redness (vs. greener for lower negative 
values), and higher positive b values increasing yellowness (vs. bluer for 
lower negative values), as calculated by the colorimeter. Primaries are 
numbered from outer to inner, so P1 is the outermost primary. 

Feathers of centre of forecrown through nape with prominent 
pointed blackish shaft-streaks up to c. 5 mm wide, each streak bordered 
on both sides by spots of Pale Pinkish Buff (121D; Munsell 7.5YR 8/4), 
distalmost spots up to c.3mm in diameter, more proximal and 
semiconcealed spots up to c.4mm,; streaks on hindcrown and nape 
narrowest. Base colour of forecrown through nape Cinnamon-Brown 
(33; L=39.0+8.1, a=8.1+0.4, b=19.2+1.3), finely vermiculated 
blackish overall. Supercilia short and not very conspicuous, extending 
only from base of eartufts to above bill, formed of small feathers that 
are whitish to pale buffy on outer webs and parts of inner webs, most 
with black tips and/or edges. Eartufts prominent, longest feather 
c. 29 mm long and c. 9 mm wide, feathers fresh, each strongly tapering 


F.R. Lambert & P. C. Rasmussen 209 Bull. B.O.C. 1998 118(4) 


toward elliptical tip, and vermiculated Cinnamon-Brown on distal half 
of outer web, with a black shaft streak up to c. 3mm wide with Pale 
Pinkish Buff spots up to c.3 mm wide along the outer edge of the 
streak, inner web and medial portion of outer web mostly Pale Pinkish 
Buff (L=53.9+2.2, a=6.3+0.6, b=23.6+0.5). Rictal bristles up to 
c. 20 mm long, mainly blackish, with barred bases of Pale Pinkish Buff 
(bars up to c. 2 mm wide) alternating with Vandyke Brown (221) bars 
up to c. 2.5 mm wide; these feathers form the darkest region of the face. 
Feathers just above eye Raw Sienna (136; Munsell 5YR 5/6), feathers 
just to the rear of and below eye slightly paler, auriculars with fine, 
vague concentric bands, broader and darker laterally, pale bands of 
auriculars Pale Pinkish Buff (219) and up to c. 2 mm wide, dark bands 
Sepia (219; Munsell 5YR 3/2) and up to c.1mm wide, medial 
auriculars have extended distal barbs up to c. 4mm long, while lateral 
auriculars mostly lack extensions. Chin and throat mostly Pale Pinkish 
Buff (L=59.0+0.8, a=4.0+0.8, b=20.6+2.0), chin slightly paler, 
more caudal feathers with fine Hair Brown (119A) barring and a few 
very small Hair Brown shaft streaks. Facial disk with incomplete, 
narrow, inconspicuous blackish border, heaviest at sides of throat, and 
bordered at rear by a poorly-defined Pale Pinkish Buff band. 

Background colour and pattern of hindneck, mantle, inner scapulars, 
and back similar to that of forecrown (mantle colour, L=29.3 + 0.3, 
a=7.4+0.2, b=14.0 40.3), with a few short blackish shaft-streaks with 
prominent Pale Pinkish Buff spots on either side of shaft streaks (up to 
c.4mm wide); on hindneck spots more profuse and streaks smaller. 
Rump and uppertail coverts appear darker (L=29.2 + 0.8, a=7.2 + 0.9, 
b=13.9+1.5) due to lack of buff spots, shaft streaks are smaller and 
more obscure, and feathers are more distinctly barred dark. Outer 
scapulars have the outer webs and a smaller area on the centre of the 
inner web Pale Pinkish Buff (L=53.5+1.4, a=3.8 + 0.8, 6=19.7 + 1.1) 
with triangular blackish tips, c. 3 mm long on smallest, uppermost pale 
scapular, and c. 5 mm long on largest, lowest scapular; the two largest 
pale scapulars also have blackish shaft streaks up to c. 1.5 mm wide. 

On the lesser coverts, the vermiculated background colour appears 
similar to but plainer than the mantle (L=33.6+1.9, a=6.8+0.2, 
b=14.9+1.1) due to lack of large pale buff spots, but most feathers 
have narrow (c. 2mm wide) Clay Color (123B; 7.5YR 7/8) bands and 
c.1mm blackish shaft streaks. The base colour of all upperwing 
coverts is similar (greater coverts: L=28.9+ 1.0, a=6.2 + 0.3, b=10.6+ 
0.6) but the markings are progressively heavier distally, and there are 
large (c.5mm) Pale Pinkish Buff (L=51.2+0.8, a=4.5+0.7, 
b=17.2 +0.6) spots to either side of the shaft streaks of the greater 
coverts. The tertials are relatively paler (L=41.441.4, a=5.940.4, 
6=14.6 + 1.2) than rest of upperparts, irregularly and vaguely banded, 
and heavily vermiculated. 

Banding of the secondaries is more definite and regular, with most 
darker bands c.5.5mm wide and Cinnamon-Brown (L=37.4 + 1.0, 
a=6.4+0.5, b=15.0+1.2) and paler bands c. 2.5 mm wide and Sayal 
Brown (223C, Munsell 7.5YR 6/6). The banding of the upper surfaces 
of the primaries is better defined, most of the dark bands on the outer 


F. R. Lambert & P. C. Rasmussen 210 Bull. B.O.C. 1998 118(4) 


webs of P3 are c.7mm wide and Raw Umber (223; L=31.2 £1.3, 
a=6.0 + 0.7, b=12.4 + 0.7) and the pale bands (of which there are 9.5) 
are c. 5.5 mm wide, and darker than Pale Pinkish Buff (L=47.2 1.9, 
a=8.9+0.2, 6=21.4+1.0). The bands are narrower on P1, the dark 
bands c.5mm wide (L=26.7+ 3.6, a=6.5+0.6, b=11.6+1.9), the 
pale bands c. 3 mm wide, and there are 8.5 pale bands on the outer web 
of P1. Most of the dark bands are finely outlined by narrow darker 
bands, the pale bands typically narrowly abut the shaft, and pale bands 
are vague on the inner webs. The tips of the primaries are paler, 
scarcely banded, and very finely vermiculated, with a dark shaft streak. 

On the underwing, the secondary coverts are Pale Pinkish Buff, the 
proximal primary coverts vermiculated brown, and the distal primary 
coverts Vandyke Brown (221) with Pale Pinkish Buff distal halves, 
forming a distinct dark patch near the bend of the wing. The under- 
surfaces of the primaries are Glaucous (79; L=37.6+0.1, a=3.9+40.2, 
b=9.7 + 0.2) with Pale Pinkish Buff bands. The axillaries are paler than 
Pale Pinkish Buff, with dark brown subterminal spots c. 3 mm long. 

The overall aspect of the underparts below the throat is very finely 
vermiculated dull brown, with the breast darker, and with prominent 
but mostly narrow blackish shaft streaks. The background colour of the 
breast is closest to Mikado Brown (121C; L=42.94+2.9, a=7.7+1.1, 
b=19.8 + 2.4), while the background colour of the lower underparts is 
distinctly paler (L=56.5 + 2.4, a=4.7 + 0.4, b=18.7 + 0.8). Each major 
feather of the underparts has a long dark shaft streak up to c. 3.5 mm 
wide but usually much narrower, two bands of Cinnamon-Drab (219C) 
finely outlined with blackish stippling, one terminal band up to c. 5 mm 
wide, separated from the other Cinnamon-Drab band (up to c. 3.5 mm 
wide) by a whitish band (L=69.0 + 2.1, a=3.2 + 0.6, b=17.2 +1.1) up 
to c. 2.5 mm wide. More proximally, there is a wider, usually concealed 
whitish band, and a pale rufous band (L=54.0+2.5, a=9.3+1.2, 
b=26.4 + 1.2). The undertail coverts and feathers around the vent are 
relatively uniform and paler than Pale Pinkish Buff, each of the longer 
feathers with an irregular dark brownish subterminal mark. 

The uppertail surface is very irregularly but rather prominently 
banded, with most dark bands c.5 mm wide (L=35.2 +0.4, a=5.2+ 
0.1, b=11.4+0.1) and pale Tawny Olive (223D) bands c. 3 mm wide. 

The tarsi are densely feathered to just above the distal joint, the 
feathering is complete around rear of tarsus, most being unmarked and 
paler than Yellow Ocher (123C; L=62.3+ 1.0, a=5.340.4, b=26.24 
0.5), the upper side with sparse short fine dark brown streaks. The toes 
and claws are relatively small and weak, and the claws have pale bases 
and dark tips. 

Specimens. 1) Holotype. 2-4) Paratypes: 2) National Museum of 
Natural History, Leiden (NNM), RMNH Kat. No. 16 (listed by 
Sharpe 1875 as manadensis No. 15, but this number is crossed out on 
the label), adult female (according to the label but a male according to 
Schlegel 1873), from “‘Sangir-Isl.’’, collected 10 January 1866 by C. P. 
Hoedt; 3) Staatliches Museum fiir Tierkunde, Dresden (MTD) No. 
C2446, adult of unknown sex, from Tabukan, collected in 1871 by 
A. B. Meyer, No. 8254 [although Meyer (1879) acknowledged failing 


Bull. B.O.C. 1998 118(4) 


211 


F. R. Lambert & P. C. Rasmussen 


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F. R. Lambert & P. C. Rasmussen 212 Bull. B.O.C. 1998 118(4) 


to properly label some specimens collected on the trip on which this 
specimen was collected, MTD C2446 closely matches the other 
Sangihe specimens in plumage and measurements]; 4) RMNH No. 
84653, adult male, from Manganitu, collected 22 May 1985 by F. G. 
and C. M. Rozendaal, skull completely ossified, weight 76 g. 

Another female specimen collected on Sangihe by Platen on 24 
January 1887 (specimen “‘b”’ of Blasius 1888) is no longer in the 
collection of the Staatliches Naturhistorisches Museum, Braunschweig, 
and is presumed to have been lost near the end of World War II (G. 
Boenigk, im litt. 1998). Since measurements given for the holotype 
(specimen ‘“‘a’’) by Blasius (1888) are very similar to those obtained 
by PCR for the same specimen, it is assumed here that Blasius’s 
measurements of the missing specimen “‘b” are comparable, and thus 
they are presented in Table 1, but they were not used in the summary 
statistics or analyses. The missing specimen was said to be redder and 
darker overall than the holotype (Blasius 1888). 

Measurements of the holotype (lengths in mm). Wing (flattened and 
straightened) 160; tail 78.0; tarsus 27.7; bill (from skull) 19.6; bill (from 
anterior edge of cere) 12.8. For additional measurements of holotype 
and those of paratypes and related species, see Table 1. 

Soft-part colours. For the holotype and Platen’s specimen “‘b’’ 
(Blasius 1888), the iris was “‘hell-gelb’’ (light yellow), and the bill and 
feet “braun’’ (brown). RMNH 84653 had “eyes bright yellow; bill 
horn; legs horn’’. An individual tape-recorded and photographed at 
night (with a flash) by FRL (Plate 5) showed pale yellow irides and a 
narrow dark rim around the eye, visible part of bill brownish-horn with 
a blackish tip, pale brownish toes, and pale-based, dark-tipped claws. 
The live specimen photographed by J. Wardill had a brownish-horn 
bill, greyish uppersurfaces to toes, soles of toes pale pinkish, and claws 
with pale bases and dark tips, but its eyes were closed in the 
photographs we have examined. 

Voice. Among related taxa for which vocalisations are known, the 
voice of collari resembles only that of manadensis, in that both are 
whistled. However, the quality, form, pitch, duration, and rhythm 
all differ greatly between the two. The songs of manadensis have 
been documented from various parts of Sulawesi. The typical song 
of manadensis is a breathy, oft-repeated whistle of two short 
staccato, syncopated elements, ‘“‘ploe- ek—ploe- ek’? or “‘oe-ek—oe-ek”’ 
(Coomans de Ruiter & Maurenbrecher 1948; Fig. 2), also rendered as 
‘“‘tona-as’’, “‘hooit”, and ‘‘dojot’? (Coomans de Ruiter 1950). Each 
phrase, which falls between c. 1-1.5 kHz, is c.0.4 sec in duration, 
commences at the lowest frequency, has a distinct frequency shift at the 
start of the second element with no decrease in amplitude, and 
terminates abruptly at the highest frequency. Other songs or calls 
reported for manadensis include a clear “‘kéte’’ or ‘“‘kiek’’, sometimes 
changing into a rapid “‘kok, kok, kok’’ or “‘kiek, kiek, kiek’’ of increasing 
amplitude but steady frequency. Alternating series of song or call 
types may also be given, ‘‘oe-ek-oe-ek-oe-ek .. . 01, 01, O1, 01, 01, OL...” 
(Coomans de Ruiter & Maurenbrecher 1948, Coomans de Ruiter 


1959). 


F.R. Lambert & P. C. Rasmussen 213 Bull. B.O.C. 1998 118(4) 


In contrast, the vocalisations of collari are strikingly different. Its 
whistled song (Fig. 2) is longer, higher-pitched, much sweeter, clearer, 
more modulated, and slurred. Each phrase is of c. 0.7 sec duration, 
consists of only one perceptible element, and falls between 
c. 1.65-1.85 kHz. A descending “‘kleeeeer’’, it commences at the highest 
frequency, is downslurred for most of its duration, and is at maximum 
amplitude during the middle two-thirds of the element. In a 
tape-recorded sequence of 11 very similar ‘“‘kleeeeer’’ phrases (Fig. 2; 
FRL), the intervals between phrases ranged from c. 8-15 sec, mean 
11.2 +2.2 sec, n=9 intervals (one interval of 28 sec was not included in 
this computation because a more distant bird was cailing; Fig. 2, 
lower). 

The latter song type (Fig. 2, lower) sounds three-noted because each 
phrase is much more variable in pitch. These phrases, which are 
separated from one another by only c. 0.3 sec, commence at c. 1.8 kHz, 
increase in frequency to c.2.0kHz, and then abruptly decrease to 
c. 1.75 kHz, remaining at that frequency for the duration. They are of 
the same sweet, liquid quality as the single-noted ‘“‘kleeeer’’ song. 

In another, more distant sequence (recording by J. Riley, NSA: 
wal997/54:4b; 2010h, 18 November 1996, Sahengbalira), six near- 
identical “‘kleeeeer’’ notes were given in 53 sec, each commencing at its 
highest pitch and being slightly downslurred (frequency range 
1.6-1.8 kHz). The intervals between phrases ranged from c. 9-11 sec, 
mean 9.4+0.9sec, n=5 intervals, and each phrase was c. 0.8 sec in 
duration. Thus, Riley’s tape-recorded sequence does not support his 
(1997b) contention that vocalisations of Sangihe birds are like those of 
manadensis, but instead it is very like the tape-recording made by FRL, 
and we know of no approach between any of the calls of the two species. 

Etymology. This species is named after our friend and colleague Dr 
Nigel J. Collar in recognition of his numerous contributions to the 
important field of bird conservation. His work has stimulated enormous 
interest in threatened birds, and has encouraged a conservation ethic 
and philosophy amongst a generation of amateur and professional 
ornithologists and birdwatchers. 


Remarks 

Differences between Otus collari and other taxa in manadensis 
superspecies 

Geographically distant and morphologically distinct taxa considered 
separate species by Rasmussen (1998: Seychelles Scops Owl Otus 
insularis and Biak Scops Owl O. beccarii) and those from islands off 
Africa are excluded from the following comparisons, as are the clearly 
different taxa of Elegant Scops Owl Otus elegans. The present 
taxonomic placement of certain other taxa is dubious, and work in 
progress is targeted toward resolving these problems, in particular the 
status of staoensis (Rasmussen & Lambert, unpubl. data). 


East Indian Ocean taxa 
From the three “East Indian Ocean”’ island endemics as a group 
(including Simeulue Scops Owl O. umbra, n=2 adult specimens 


F. R. Lambert & P. C. Rasmussen 214 Bull. B.O.C. 1998 118(4) 


examined; Enggano Scops Owl O. enganensis, n=4; and Nicobar Scops 
Owl O. alius, n=2, Rasmussen 1998; but excluding the Mentawai 
Scops Owl Otus mentawz), collari differs in being more streaked overall, 
having longer eartufts, a more prominent facial disk, and more spotted 
upperparts. However, collari and siaoensis are the only Wallacean taxa of 
this superspecies with such narrow wings as the East Indian Ocean 
group, but both of the former taxa differ from the latter group in that the 
second and third primaries (from outermost) are shorter, which gives 
them more pointed wingtips (Table 1). Additionally, collari differs from 
alius (see plate in Rasmussen 1998) in that its supercilium is whiter and 
more prominent, in overall plumage it is much more heavily streaked, 
with spotting rather than barring above, its pale scapular spots are larger 
with triangular black tips, its tarsi are feathered more distally, and its feet 
and claws are smaller. From enganensis, collari differs in its smaller size, 
especially its bill, feet and claws, in lacking long extensions of the distal 
barbs of the rear auriculars, in having larger, blacker scapular tips, and in 
being much less uniform and less rufous overall. From umbra, collari 
differs in its much longer wings and tail and much less uniform, less 
rufous overall plumage. 


Taxa from outlying islands in Sulawesi region 

Otus [m.] staoensis, despite its provenance, differs more strongly from 
manadensis and collari than the latter two do from each other (Plate 4, 
Table 1). Nevertheless, szaoensis and collari appear to share a pattern of 
shortfalls of primaries from the wingpoint that differs from that of all 
the Moluccan and Lesser Sundas taxa of magicus, as well as manadensis, 
the following three taxa from islands to the east of Sulawesi, and the 
more distantly related beccarit and insularis (Rasmussen 1998). The 
apparent similarity in wing shape notwithstanding, collam differs 
greatly from szaoensis in numerous characters. The former has much 
longer wings and tail, although the bill, feet and claws are of 
approximately the same size, its eartufts are longer, less full and more 
pointed, its throat is more heavily marked and less conspicuously pale, 
it lacks a prominent pale nuchal collar, its underparts pattern is much 
more vermiculated, more regularly barred, and drabber in colour, with 
the black shaft-streaking greatly predominating over the barring, its 
remiges and rectrices have fewer bands, which are outlined on both 
edges by narrow blackish bands, and its tail is much more broadly, 
irregularly, and less strongly banded. 

Compared to Otus [m.] sulaensis (Finsch 1898, King 1997; Sula 
Islands, east of Sulawesi, n=2 adults, entire world holdings), collari is 
smaller, less heavily marked, and paler overall (Table 1). Its auriculars 
are paler, its eartufts blotchier and unbarred, its upperparts more broadly 
streaked, its scapular spots lack large blackish blotches, the pale bands in 
its primaries are larger and extend to the shaft and to some extent the 
inner web, its tail is more prominently banded, and its tarsi are much 
more extensively feathered, especially on the rear (plantar) surface. 

Otus collari differs from O. [m.] Ralidupae (Tukangbesi, south-east of 
Sulawesi, n=2 adults, entire world holdings) in being smaller, darker, 
and browner overall (Table 1). Its eartufts are also much more heavily 


F.R. Lambert & P. C. Rasmussen 215 Bull. B.O.C. 1998 118(4) 


marked, its scapular spots have large black terminal triangles and lack 
fine black transverse lines, its underparts have more distinct, finer 
barring, the upper surfaces of its outer primaries have much wider and 
more uniformly dark bands, but the undersides of the inner webs of the 
primaries have less prominent pale barring, and its tertials and upper 
tail surface are more prominently banded. 

From O [m.] mendeni (Peleng Island, off eastern Sulawesi, n=3 
adults, entire world holdings), collari differs greatly in its much longer 
wing and tail. Additionally, its eartufts are more heavily streaked and 
spotted, it has a paler bill, the darkest area of its face is between the bill 
and eyes, its upperparts have much broader streaks and larger pale 
spots, its overall plumage is much more uniformly brown, the streaking 
on its underparts is narrower, less blotchy, and lacks rufous areas 
around the black shaft streaks, its scapular spots have larger pale areas 
(rather than having black central cross-lines) and larger triangular black 
tips, its tertials are more prominently banded, and its tarsus is feathered 
much more distally. 


Otus magicus 

Compared to O. magicus albiventris and O. m. tempestatis, the small 
Lesser Sundas (Nusa Tenggara) subspecies of Otus magicus, collari 
differs in that the auriculars and area around the eye do not form a 
distinct dark patch, its white supercilium does not extend to the front of 
the face, its bill is browner, its upperparts are more twin-spotted, its 
lower underparts have smaller, less conspicuous white markings, its 
primaries have broader dark bands, its tertials are less prominently and 
less evenly banded, and its tail is much more broadly but less distinctly 
dark-banded. Otus collari also differs from albiventris in that its eartufts 
are shorter, less pointed, more blotched, and have buffer bases, and its 
underparts are more finely vermiculated with fewer heavy black streaks. 
Additionally, collar: differs from tempestatis in that its pale scapular 
spots have larger black triangular tips and lack narrow blackish cross 
marks. The five Moluccan forms of Otus magicus (Table 1) are all 
considerably larger than collari, and none closely resemble it in 
plumage. 


Philippine taxa 

To the north of Sangihe, the nearest related taxa are four disjunct 
Philippine races of Mantanani Scops Owl O. mantananensis: O. m. 
cuyensis (Cuyo I., west-central Philippines, n=3); O. m. romblonis 
(Romblon, Sibuyan, and nearby islets, n=5); O. m._ sibutuensis 
[including steerei, type examined), Sibutu and Tumindao, Sulu 
Archipelago, n=15]; and O. m. mantananensis (islets between Borneo 
and Palawan, n=2). Compared to all these taxa (combined), collari is 
smaller, especially its bill and feet, although its wings are longer and 
narrower (‘Table 1), the lower edge of its throat is less streaked, its facial 
disk is less distinct, and its tarsi are more extensively feathered. 
Additional differences exist between collari and each race of 
mantananensis. 


F.R. Lambert & P. C. Rasmussen 216 Bull. B.O.C. 1998 118(4) 


Habitat, ecology, and conservation 

The 1985 Otus collavi specimen was collected “in cultivation of 
coconut, nutmeg and secondary growth, along stream, c.50m.”’ at 
Manganitu (label data), and the species was also reported during the 
same time period from the upper slopes of Mt. Sahendaruman (F. G. 
Rozendaal, zm litt. 1998). FRL’s tape-recording and photographs were 
made of a bird calling 2—5 m above the ground in the understorey and 
lower parts of larger trees just before and after dusk on 30 July 1996 
above ‘Telawid Atas, at about 315 m altitude on the slopes of Mt. Awu, 
in a valley of remnant semi-natural habitat with tall durian and other 
trees, bamboo and scrub on the edge of a mixed plantation. Otus collari 
was also heard calling on 8 and 9 September at Telawid Atas (Riley 
1997a), in November and December 1996 in forest on Mt. 
Sahengbalira and from agricultural areas around Telawid (Riley 
1997b), and a captive obtained in Tahuna was said by locals to have 
come from nearby mixed plantations (Riley 1997b). Thus it appears to 
be widespread throughout Sangihe, from the coast to at least the lower 
parts of the hills. Given its apparent tolerance of mixed plantations, a 
habitat which has dominated the landscape of the island since the early 
part of the 20th Century (Heringa 1921, Whitten et al. 1987), Otus 
collart seems unlikely to be threatened. 


Acknowledgements 


For information, tapes, and other assistance we thank especially B. F. King, J. Riley, J. 
Wardill, F. G. Rozendaal, R. Ranft (National Sound Archives, London), and J. T. 
Marshall, Jr. Specimens were examined in or borrowed from the collections listed below, 
for which we thank the following: G. F. Barrowclough, J. Cracraft, M. LeCroy, P. Sweet 
and A. Andors, American Museum of Natural History (AMNH); L. Joseph, L. Bevier 
and D. Agro, Academy of Natural Sciences of Philadelphia; M. P. Walters and R. 
Prys-Jones, The Natural History Museum, Tring, U.K.; A. Rahmani and S. Unnithan, 
Bombay Natural History Society; G. Hess, Delaware Museum of Natural History; D. 
Willard, Field Museum of Natural History; R. A. Paynter, Jr. and A. Pirie, Museum of 
Comparative Zoology, Harvard University; C. and J.-F. Voisin, Museum National 
d’Histoire Naturelle, Paris; S. Eck (MTD, Dresden); R. Dekker (NNM, Leiden); G. 
Boenigk (SNMB, Braunschweig); R. B. Payne and J. Hinshaw, University of Michigan 
Museum of Zoology; R. S. Symonds, University Museum of Zoology, Cambridge; S. L. 
Olson, J. P. Angle, B. M. McPhelim and J. Dean, National Museum of Natural History 
(USNM, Washington, D.C.); S. Frahnert and J. Fiebig, Museum ftir Naturkunde, 
Berlin; and K. Roselaar, Zoologisch Museum, University of Amsterdam. The 
manuscript was improved by N. J. Collar, R. C. Banks and G. R. Graves. J. Wardill 
provided photos, and J. C. Anderton painted the accompanying original watercolour 
plate. 


References: 

Blasius, W. 1888. Die Vogel von Gross-Sanghir. Ornis 4: 527-646. 

hemene de Ruiter, L. 1950. Vogel van het Quarles-Gebergte (ZW. Centraal Celebes). 
Ardea 38: 40-182. 

Coomans de Ruiter, L. & Maurenbrecher, L. L. A. 1948. Stadsvogels van Makassar 
(Zuid-Celebes). Ardea 36: 163-198. 

Finsch, O. 1898. Uber Scops magicus (S. Mill.) und die verwandten Arten. Notes Leyden 
Mus. 20: 163-184. 

Graves, G. R. 1997. Colorimetric and morphometric gradients in Colombian populations 
of Dusky Antbirds (Cercomacra tyrannina), with a description of a new species, 
Cercomacra parkerit. Orn. Monogr. 48: 21-35. 


M. LeCroy & W.S. Peckover 217 Bull. B.O.C. 1998 118(4) 


Heringa, P. 1921. Rapport over de begroeining van de Sangi- en Talaud-eilanden. 
Tectona 14: 733-746. 

King, B. F. 1997. Checklist of the birds of Eurasia. Ibis, Vista, California. 

Lambert, F. R. 1998a. A new species of Gymnocrex rail from the Talaud Islands, 
Indonesia. Forktail 13: 1-6. 

Lambert, F. R. 1998b. A new species of Amaurornis from the Talaud Islands, Indonesia, 
and a review of taxonomy of bush hens occurring from the Philippines to 
Australasia. Bull. Brit. Orn. Cl. 118: 67-82. 

Marshall, J. T. 1978. Systematics of smaller Asian night birds based on voice. Orn. 
Monogr. No. 25: 1-58. 

Marshall, J. T. & King, B. F. 1988. Genus Otus. Pp. 331-336 in: Amadon, D. & Bull, J. 
Hawks and owls of the world: a distributional and taxonomic list. Proc. Western 
Found. Vert. Zool. 3: 295-357. 

Meyer, A. B. 1879. Field notes on the birds of Celebes. Part 1. Psittaci, Rapaces, and 
Picariae. Ibis (4th Ser.) 3: 43-70. 

Meyer, A. B. 1884. Ueber neue und ungentigend bekannte Végel, Nester und Eier aus 
dem Ostindischen Archipel im KG6nigl. Zoologischen Museum zu Dresden. Abh. 
Naturwiss. Ges. Isis Dresden 1: 1-64. 

Meyer, A. B. & Wiglesworth, I. W. 1898. The birds of Celebes and the neighbouring islands. 
R. Friedlander, Berlin. 

Munsell. 1997. Munsell Color Charts for Plant Tissues. Gretagmacbeth, New Windsor, 
New York. 

Rasmussen, P. C. 1998. A new scops owl from Great Nicobar Island. Bull. Brit. Orn. Cl. 
118: 141-153. 

Riley, J. 1997a. The birds of Sangihe and Talaud, north Sulawesi. Kukila 9: 3-36. 

Riley, J. 1997b. Biological surveys and conservation priorities on the Sangihe and Talaud 
islands, Indonesia. CBS Conservation Publications, Cambridge. 

Rozendaal, F. G. & Lambert, F. R. In press. The taxonomic and conservation status of 
Pinarolestes sanghirensis Oustalet 1881. Forktail. 

Schlegel, H. 1873. Revue de la collection des oiseaux de proie faisant partie du Musée des 
Pays-Bas. Aves Noctuae. Oti. Tome II, Monogr. 11. E. J. Brill, Leiden. 

Sharpe, R. B. 1875. Catalogue of the birds in the British Museum. Vol. II. Striges. 
Trustees of the British Museum, London. 

Smithe, F. B. 1975. Naturalist’s color guide. American Museum of Natural History, New 
York. 

Stattersfield, A. J., Crosby, M. J., Long, A. J. & Wege, D. G. 1998. Endemic bird areas of 
the world. BirdLife Conserv. Ser. No. 7. BirdLife International, Cambridge. 

White, C. M. N. & Bruce, M. D. 1986. The birds of Wallacea. B.O.U. Check-list No. 7. 
British Ornithologists’ Union, London. 

Whitten, A. J., Bishop, K. D., Nash, S. V. & Clayton, L. 1987. One or more extinctions 
from Sulawesi, Indonesia? Cons. Biol. 1: 42-48. 


Addresses: Frank R. Lambert, BirdLife International, PO Box 310/Boo, Bogor 16003, 
Indonesia; Pamela C. Rasmussen, Room 336 NHB MRC 114, Smithsonian 
Institution, Washington, D.C. 20560-0131, U.S.A. 


© British Ornithologists’ Club 1998 


Misima’s missing birds 


by Mary LeCroy & William S. Peckover 
Received 10 September 1997 


We observed 37 species of birds on Misima Island, Louisiade 
Archipelago, Papua New Guinea during a visit from 1-12 October 
1993. Comparison of our list with species collected on the same island 
by A. S. Meek in 1897 (Hartert 1898a,b, 1899) led to the discovery that 


M. LeCroy & W.S. Peckover 218 Bull. B.O.C. 1998 118(4) 


a number of species were “‘missing”’ from our list. Thinking that 
perhaps the missing species had been victims of the extensive habitat 
alterations that have taken place in the past century, we undertook a 
careful analysis of avian records from Misima. This paper presents our 
results and shows that the uncritical use of the published records might 
have led to false conclusions, but that careful analysis of archival and 
specimen records may provide unexpected discoveries and useful 
insights. 

The Louisiade Archipelago has long been known to Europeans. 
Captain Luis Baez de Torres discovered it in 1606 and named it Tierra 
de San Buenaventura (Hilder 1980: 24 and fig. 6), but he sailed south of 
the bordering reefs and islands without making a landfall on or naming 
any of the islands individually. Perhaps unaware of the earlier Spanish 
name, Captain Louis-Antoine de Bougainville in 1768 called the group 
the Louisiade Archipelago to honour King Louis XV of France. On 
14 June 1793, Captain A. J. R. Bruni D’Entrecasteaux sighted the 
island now known by its local name of Misima and named it St. Aignan 
for Frenay de Saint-Aignan, Lieutenant on the Recherche, a frigate 
under D’Entrecasteaux’s command (Wichmann 1909: 267) (Fig. 1). 


Existing collections and records 


Between 1879 and 1930 two large and several small collections of birds 
were made on Misima. The largest was made by A. S. Meek who, with 
his brother W. G. Meek, one of the Eichhorn brothers never mentioned 
by name, and local assistants, visited in 1897, collecting for Lord 
Walter Rothschild. The part of this collection reported on by Hartert 
(1898a,b, 1899) is now in the American Museum of Natural History 
(AMNH) in NewYork. Hartert gives no clear idea of the number of 
specimens or dates of collection but lists 65 species and dates between 
late July and December. Meek (1913: 89) says that he spent three 
months on Misima. We were able to trace 273 former Rothschild 
Collection specimens at AMNH, including all species listed by Hartert 
except Ducula zoeae, and three species not listed by him, Pluvzalis 
squatarola, Sterna sumatrana, and Lorius hypoinochrous. The nests and 
eggs collected remain in the Natural History Museum, ‘Tring, U.K. 
formerly the British Museum (Natural History) (BMNH); most dates 
given for these by Hartert (1899) are in November and December. 

The second largest collection was made by Hannibal Hamlin (MS), 
when leader of the AMNH Whitney South Sea Expedition, and two 
assistants. They spent 21-30 July 1930 at Bwagabwaga Bay on the 
south side of Misima, and 95 specimens of 23 species were catalogued, 
of which we traced 93. Weights and conditions of gonads are noted 
on the labels, as are some soft parts colours. This collection has not 
been reported as a whole, although specimens from it have been 
incorporated in various papers resulting from the Whitney Expedition 
and in Mayr (1941). 

Hamlin (MS) was also in the Bonvouloir Islands in 1928: on 
Hastings Island on 8 October, collecting 27 specimens of seven species 
and seeing an additional one; on East Island on 10—11 October, 


Bull. B.O.C. 1998 118(4) 


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collecting 58 specimens of 11 species. On 28-29 April 1929 and 18 
July 1930 he was on Panasesa [‘“‘Pana’”’ means “‘island’’ (Brass 1959: 
33)], Conflict Group, where he collected 29 specimens of six species 
and observed five additional species; and on 30-31 July 1930 on 
Panapompom, Deboyne Islands, where he collected 15 specimens of six 
species. On 19-20 and 30 July 1930 he was on Nivani Island, Deboyne 
Islands, where he recorded seeing nine species, apparently without 
collecting. These records have been included in Table 1. 

Tristram (1882) reported on a collection made in 1879 by G. E. 
Richards, R.N., at Blanche Bay, New Britain, and on Misima. Richards 
was captain of the Royal Navy warship Renard, involved in survey 
work in the Solomon Sea (Wichmann, 1910: 269, 1912: 828-829). He 
sent a total of 62 species from both localities, but a complete list is not 
given and only Lorius hypoinochrous, Esacus magnirostris, Pluvialis 
fulva, and Ducula pinon salvadoriu (Carpodacus salvadori, described as 
new) were definitely ascribed to Misima. 

Other small collections made about the same time were reported on 
by DeVis (1890, 1892) and Tristram (1889b). The specimens listed by 
DeVis were from collections made for Sir William MacGregor, 
Administrator of British New Guinea, during his official visits to the 
Louisiade Archipelago: the first, 11 species from Misima, was made 
during his first ten months in office (September 1888 through June 
1889), but no dates are given by DeVis; the second, during June and 
July 1891, when collections were made on Maisima (four species), 
nearby “Renard Island’? (=Kimuta Island, nine species) and 
Panapompom (one species). We have considered only the species that 
were listed with specific localities (Table 1). Some specimens from the 
latter of these two collections are in the Queensland Museum (QM) in 
Brisbane. 

Tristram (1889b) reported on the birds collected by Basil H. 
Thomson, Secretary to Sir William MacGregor, while on a 
government exploring trip. He spent 20-23 October 1888 on the island, 
anchoring “‘on the lee side at a spot which would be quite unsafe in the 
north-west monsoon’’. His group, including geologists, apparently 
walked across the centre of the island. ‘“We found that the limestone 
hills which compose the centre of the island were honey-combed with 
caves and densely timbered. We crossed the range and descended to a 
most romantic spot called by the natives Kaiaba .. . The eastern part of 
the island consists of very rugged hills, through which the streams have 
cut very deep and narrow gorges’’ (Thomson, 1889). Tristram listed 
five species from Misima, including Cinnyris christianae (= Nectarinia 
aspasia christianae) which he described as new. 

Maitland (1893), who was unable to pinpoint Kaiaba, explored the 
headwaters of Gulewa Creek and thought perhaps this was the same 
area described by Thomson, implying that ‘Thomson crossed from 
south to north. More recently, the Fifth Archbold Expedition spent 
14 July—14 August 1957 on Misima. Brass (1959: 34) notes that a visit 
was made to “‘a big cave a mile or more up a creek from Ehaus [= Eiaus] 
village, about 4 miles west of camp. Caves on three levels were 
reported. Only the middle-level cave, seemingly called Kuiaba, was 


Bull. B.O.C. 1998 118(4) 


221 


M. LeCroy & W.S. Peckover 


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entered, in company with villagers from Ehaus.’’ We feel that Thomson 
and his group probably anchored on the north coast, certainly ‘‘unsafe 
in the north-west monsoon’’, and perhaps not far from our locality of 
Gulewa where a track across the island, now a road, has its northern 
terminus. If they then walked across on this track they would come out 
on the south coast not far from Eiaus and Kuiaba cave. Additional 
circumstantial evidence that ‘Thomson landed on the north coast is the 
establishment in 1889 of government headquarters for the Louisiades at 
Siagara on the north coast, where it remained only briefly before being 
moved to the protected site of Bwagaoia (Brass 1959: 19). It may be 
that Thomson’ 1888 visit affected the choice of the first government 
station. If we are correct in our interpretation of Thomson’s route, 
the “densely timbered’’ hills that he reported are largely denuded of 
forest today. 

Brian Finch (1985) added two species to the Misima list from 
observations at the Misima airstrip at Bwagaoia in February 1985: 
Glareola maldivarum and Calidris ferruginea. More recently, Len 
Tolhurst (1996: 34-35) has reported two records for Misima: a 
breeding Amaurornis olivaceus and Merops ornatus. However, Merops 
ornatus was collected earlier by Meek (Hartert 1899: 212) and also by 
Hamlin (unpublished). No collections or observations have been made 
on Mt. Diatau (1,038 m) and Mt. Apatikaiogean (703 m), the two 
highest points on the island. Meek (1913: 86) mentions that his 
collectors obtained butterflies at higher altitudes, even though his base 
camp was at Bugoiya Harbour, and it is possible that some of his birds 
‘came from these areas as well. 


Misima in 1993 


Most of our stay on Misima was spent at and near the village of 
Gulewa, in a mixed forest—garden area ca. 200 m above sea level, 2 km 
south of the village. We spent 1-3 and 12 October at Misima Mines 
Pty. Ltd. headquarters and some observations were made there, and 
between there and Bwagaoia. All of these records are listed in Table 1. 

At the time of our visit, Misima had had no rain in five months. ‘The 
hillsides were very dry and local people were cutting and burning large 
tracts of the remaining forest for gardens. Many of the slopes that were 
being denuded of forest were very steep (up to 60°) and will be prone to 
erosion. Large trees were felled and burned on the site. Smaller timber 
was utilized to help prevent erosion, to build pig-proof fences around 
the gardens, or to build houses. Light showers early in our stay became 
more numerous and heavier, until by the time we left Gulewa the fires 
were all out and the air clear. Gardens already extended to the tops of 
some of the hills, although higher hills were still topped by forest. ‘True 
gullies were not cultivated and some of these had sago trees and tree 
ferns growing in them. We did not see tree ferns or palm trees in the 
forest, but there were a few low palmetto-like plants on the forest floor. 

Misima Mines Pty. Ltd. has provided numerous amenities to the 
local people, including a good road around the eastern half of the island 
and piped water into most villages. Most small streams near the coast 


M. LeCroy & W.S. Peckover 225 Bull. B.O.C. 1998 118(4) 


were dry, but water was still available at the village faucets. Larger 
rivers were still running and people used them for bathing and for 
washing clothes and dishes. 

The country between the mine and Bwagaoia was mostly coconut 
plantations, with cocoa planted beneath the coconut trees. Both were 
old, unkempt, and dying; there were very few new plantings. There 
were some banana plantations in lower, presumably wetter, locations. 
Much of the northeastern end of the island is high, with cliffs, often 
deeply undercut by waves on the windward side. On the north coast, 
just outside Bwagaoia, there is an area of dry eucalypt forest. The 
beaches around Gulewa are coralline rock, sharp and pitted, with few 
areas of sand. 

The forest remaining near the coast was very disturbed and largely 
dry, but there was some lush sago forest, even though the ground in it 
was dry at this season. Trails into the gardens from Gulewa were 
everywhere steep, and the soil in the gardens was very dry and rocky. 
Several residents were pasturing cattle in scrubby land. 

We spent 4 days at a garden house about 200 m above Gulewa. The 
walk to this place began in a dry area of planted sago, coconut, 
breadfruit, cocoa, and betel nut. A few big forest trees had also been 
left in place, including “‘chestnut’”’, Terminalia sp. As we started to 
climb, we went through established garden patches and then into 
newly cut forest, some still smouldering. This part was very steep and 
rocky. 

Forest, apparently original, covered a small area adjacent to the 
garden house and topping the hill. Here there were tall buttressed trees, 
25-30 m high, and the ground under them was relatively free of 
undergrowth. The interior of this forest was quite damp and everything 
green. Most mornings there was a heavy dew; this probably also gave 
the gardens some moisture, even though the soil was dry and powdery. 

Human population growth in Misima is very rapid and this is 
undoubtedly a factor in the wholesale forest destruction. Catapults are 
much in evidence, including very large ones, 2—2.5 m long with a “Y”’ 
at the top about 30 cm deep. The basal end of the stock is pointed and 
is stuck in the ground and slanted to aim the catapult. Flying foxes and 
probably large birds are shot in this way. 

Misima conforms to the pattern of bird distribution on other oceanic 
islands, with relatively few species of birds, but most of these common, 
and few understorey birds. 


Re-analysis of Meek’s collection 


When we compared our sight records with the species known from 
Misima, it became apparent that Meek was the only collector to have 
obtained some species. Further analysis indicated that these were either 
very widespread species, migrants or species that are small island 
specialists, and we began to suspect that not all of his specimens came 
from Misima, even though they were labelled “‘St. Aignan’’ and Hartert 
had apparently accepted this locality. Our analysis also showed that 
he had collected bird specimens from 31 July—11 September and 


M. LeCroy & W.S. Peckover 226 Bull. B.O.C. 1998 118(4) 


26 November—28 December. Most of the anomalous species were taken 
in late November and December, although a few were taken on 31 July 
and 5-11 September. Fortunately, we were able to refer to Meek’s 
letters to Rothschild, Hartert or Jordan at Tring, which are now in the 
Library of BMNH. One of us (WSP) has indexed these letters and they 
are now on microfilm. Microfilm letter nos 55-64 are the ones 
applicable to his Misima trip. 

Letter no. 55, 9 July 1897, Samarai, Meek to Hartert: “... I shall 
draw £50 against them [his last collection] for current expenses until | 
can get a large consignment from St. Aignans, where I hope to be going 
in about a week from date. ‘The best way I think will be for you to let 
_Gerrard see what you select and for him to value them. I send them to 
you as I wish (after seeing your letter of 20th April) you to understand 
that I wish to be as fair as lay [szc.] in my power to be... 

Letter no. 56, 24 July 1897, Cutter Calliope [Meek’s boat], Samarai, 
Meek to Hartert: “... Am expecting to get away tomorrow or day after 
for St. Aignans... I shall send a consignment from St. Aignans as soon 
as I can conscientiously do so, for as you’ve no doubt guessed by now, 
I’m rather short of money ... Most probably there will be two 
collections from St. Aignans...”’ 

Letter no. 57, 19 August 1897, St. Aignans I., British New Guinea, 
Meek to Rothschild: “It [the Misima birdwing butterfly] is very 
striking after being accustomed to the green insect; which I have now 
taken or seen, from Cape Vogel on mainland to the Engineer Group 
(the nearest group of islands west of St. Aignan’s excepting the 
Conflicts). At the Engineer Group, the females run nearly black, nearly 
all having black forewings. I captured eight there on my way here.”’ 

Another page in the same letter, dated 29 August, “... at present I 
have something over two hundred bird skins...” 

Letter no. 58, 3 September, St. Aignan’s I., Meek to Jordan: “... 
We have skinned about 230 birds ... so far I have only forty six species 
of bird, that I can recollect without referring to my books...” 

Letter no. 59, 10 October 1897, St. Aignan I., Meek to Hartert: 
‘Having had scarcely anything but bad news to communicate up to the 
present time, you will be pleased to know that fortune has taken a 
favourable turn and am now happy in being able to say that I have now 
a very fair collection to [szc.] little over two months work . . . This place 
is extremely poor as regards birds and mammals. I shall send 
collections by first opportunity ...”’ 

Same letter, but dated 21 November: “... Shall be sending 
collections within a fortnight; seven cases in all...” 

Same letter, but dated 29 November, Samarai: “... Am sending six 
cases of Nat. History specimens and am drawing £200 against them. I 
think you will be pleased with them. Be on Rossell Island early in 
January. Written in haste.”’ 

Letter no. 60, 18 October 1897, St. Aignan’s I., Meek to Rothschild: 
‘<’.. When these collections are dispatched, I intend to sail down to 
Rossell Island, leaving my collectors at a small island close here called 
Kimuta, where the Ornithoptera [bird-wing butterflies] are not quite as 


> 


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M. LeCroy & W. S. Peckover 227 Bull. B.O.C. 1998 118(4) 


Letters nos 61 and 62 have no year and are out of order. They are 
letters nos 45 and 46 in chronological order, having been written in 
1896. They are not concerned with either Misima or Rossel. 

Letter no. 63, 10 January 1897, Rossel I., Meek to Jordan: “... 
Arrived here two days ago from Misima (St. Aignans’) bringing my 
men and five or six tons of yams. This last month I have been cruising 
around the islands, getting yams and boys. I left my boys on a small 
island off St. Aignan’s named Kimuta. They got three more specimens 
of Ornithoptera identical with St. Aignan’s and a good quantity of bird 
ewias .. 

Letter no. 64, 11 January 1897, Rossel I., Meek to Rothschild: “... I 
left my men on a small island SE of St. Aignan’s named Kimuta. They 
managed to get three more males of the blue Ornithoptera and some 
two hundred bird skins ...”’ 

The 273 specimens from the Rothschild Collection, and one that 
came to AMNH from J. H. Fleming of ‘Toronto and was never a part of 
the Rothschild Collection, did not seem to account for the number of 
birds that Meek referred to in the letters quoted above. When we listed 
AMNH specimens in field number order, we found that the lowest was 
721, collected on 31 July, and the highest was 1233, collected on 
28 December. This gives a total of 513, of which Rothschild selected 
only 273. We discovered this discrepancy to be explained by the fact 
that Rothschild frequently purchased only a portion of a collection, 
turning over the remainder to a dealer to sell for Meek. This is alluded 
to in the reference to Gerrard (E. Gerrard & Sons, a London dealer) 
in letter no.55 above. Miriam Rothschild (1983: 158) states that 
Rothschild had an agreement with Meek to take “Six specimens of 
every species at 6/6 each, with a bonus of £4 for every new species’’. 
When one looks at the entire collection listed in field number order, it 
is apparent that Meek collected long series of common birds on certain 
days, interspersing that with the fortuitous collecting of rarer species 
when they could be obtained. Rothschild selected a series of six 
specimens of the common species and apparently took all specimens of 
species represented by fewer than six. 

Fleming’s collection is now in the Royal Ontario Museum (ROM) 
and includes 74 specimens from “St. Aignan’”’, purchased by Fleming 
from Gerrard. These have not added any species to those reported by 
Hartert. Some of the Fleming specimens of Pachycephala pectoralis and 
Zosterops griseotincta are labelled ‘‘cotype’” (=syntype in today’s 
terminology). Their standing depends on whether or not Hartert wrote 
the 1899 paper before the collection was broken up. Most of the time he 
did not list the exact number of specimens, and internal evidence is 
rarely helpful. However, two bits of evidence indicate that he only 
considered the part of the collection that Rothschild bought, resulting 
in Fleming’s specimens have no nomenclatural standing. 

In his discussion of Monarcha trivirgatus, Hartert says: “‘Of this rare 
bird Meek sent four males and one female (italics his) from St. Aignan’’. 
The AMNH has three adult males, one adult female (identical in 
appearance to the males) and one female in immature plumage. The 
ROM has three adult males, two adult females and one immature 


M. LeCroy & W.S. Peckover 228 Bull. B.O.C. 1998 118(4) 


female; and the BMNH has three adult males and an adult female, none 
of which is mentioned by Hartert. Had he had all of these specimens, 
he would surely not have considered it rare. 

Hartert gives measurements of two adult males of Myzagra rubecula 
saying, “‘the adult males (italics his) are giants ...’? AMNH has two 
adult males, one male in late immature plumage, two females, and one 
female?; the ROM has a further three adult males, one immature male, 
and two adult females; and the BMNH has an adult male and female. 
Had Hartert had them, he would surely have measured the additional 
““giant’’ males. 

Replies to our enquiries disclosed an additional 48 Meek St. Aignan 
specimens at the BMNH, two at the National Museum of Natural 
History, Washington, DC (USNM), and one at the Liverpool Museum 
(LIVCM). None of these changed the species tally but did slightly alter 
dates, and they have been included in the preparation and discussion of 
‘Table 1. Also, the field numbers on non-AMNH specimens extended 
the collection numbers from 720 to 1234, giving a total of 515 birdskins 
originally collected, of which we traced 398. 

One of the specimens at the BMNH came to them via the Hewitt 
Collection, the two at the USNM came from the Swales Collection, 
and the one at LIVCM was from the Barrow Collection. These 
specimens, once part of large private collections, lead us to believe that 
Meek specimens were acquired from Gerrard by many collectors and 
more may be extant but widely scattered. We would appreciate hearing 
of any additional Meek St. Aignan (Misima) specimens that are held in 
other museums. 


Collecting localities and dates, a discussion of Table 1 


We know that Meek was on Basilaki I. on 25 July (see account of 
Ducula zoeae, Meek no. 719, below) and on Misima by 2 August. No 
specimens were collected on 1 August. In letter no. 57, quoted above, 
Meek mentions that he collected the birdwing butterfly Trozdes priamus 
in the Engineer Group on his way to Misima, so we know that he made 
at least one stop on small islands between 25 July and 2 August; 
unfortunately the butterfly specimens have no exact date. 

Six species of birds were collected on 31 July (ten specimens, 
beginning with Meek no. 720, one of which was not traced): Columba 
vitiensis, Ptilinopus rivoli, Monarcha guttula, Pachycephala pectoralis, 
Zosterops griseotinctus, and Myzomela albigula. 'The last three are well 
known as species occurring on small islands and have not been seen or 
collected by others on Misima, except for a specimen of “‘Pachycephala 
melanura’ reported by DeVis (1892: 96) which cannot now be found in 
the Queensland Museum; until corroborated, it should be removed 
from the Misima list. Columba vitiensis is widespread but never 
common, and while frequently collected on small islands, this is not 
always the case. It was collected on Misima by Hamlin. Ptilinopus rivoli 
and Monarcha guttula are discussed below. In Table 1, 31 July birds 
are listed separately and are considered to be from one of the groups of 
small islands between Basilaki and Misima, probably the Engineer 


M. LeCroy & W.S. Peckover 229 Bull. B.O.C. 1998 118(4) 


Group; in any case, they do not change the species lists for Misima 
and Kimuta. During August, none of the small island species was 
collected. 

From careful analysis of all the records, we think it likely that Meek 
moved some collectors from Misima to Kimuta early in September and 
had collectors in both localities, during at least part of the period 
between 5-11 September. Certainly, by 8 September he had some 
collectors on Kimuta. The only specimens in the collection with a 
locality other than “St. Aignan”’ are four specimens of Aplonis 
cantoroides collected on 8 September and marked ‘“‘small island off 
St. Aignan’”’ which, from letters nos 60 and 63 above, must refer to 
Kimuta. 

The ten species collected on 1-4 September are included in the 
Misima list in Table 1 as they are widespread, have been reported by 
others on Misima, and all except Collocalia vanikorensis were also 
collected by Meek in August. 

Specimens of the 27 species collected 5-11 September have been 
listed separately in Table 1. Only three of these were otherwise 
collected by Meek on both islands: Tringa hypoleucos, Ptilinopus rivolit, 
and Monarcha trivirgatus. Four species were collected only during 
those dates: Anas superciliosa, Ducula pistrinaria, Ducula bicolor, and 
Eudynamis taitensis. Ducula bicolor is reported by Tristram (1889b: 558) 
from Misima; the other three species are not otherwise reported but 
might be expected on either island. These three species are not counted 
for either island. 

Eleven species were collected by Meek both during 5—11 September 
and on Misima in August: Megapodius freycinet, Macropygia 
amboinensis, Ptilinopus superbus, Micropsitia pusio, Geoffroyus geoffroyt, 
Merops ornatus, Gerygone magnirostris, Rhipidura rufiventris, Monarcha 
guitula, Nectarinia aspasia, and Aplonis metallica. Eight of these are 
known only from Muisima; Merops ornatus, Monarcha guttula and 
Nectarinia aspasia have been reported from other small islands, but not 
from Kimuta (Table 1). 

On the other hand, nine species were collected during 5-11 
September and also on Kimuta in November and December: Pluvialis 
fulva, Chalcophaps indica, Trichoglossus haematodus, Halcyon chloris, 
Rhipidura rufifrons, Monarcha cinerascens, Pachycephala pectoralis, 
Zosterops griseotincta, and Aplonis cantoroides. Five of these have not 
been seen or collected on Misima. Pluvialis fulva has been collected by 
others, and we saw Chalcophaps indica, Halcyon chloris and Aplonis 
cantoroides (‘Table 1). Specimens collected on Kimuta on 5-11 
September may have convinced Meek that it would be productive to 
leave collectors on Kimuta in late November. 

Meek’s activities between the last half of September and the end of 
November are not documented, but we are sure that he was not 
collecting birds because the highest specimen number from 11 
September is 1030 and the lowest from 26 November is 1031. 
Apparently he was concentrating on collecting insects and other 
animals, as he was still writing from “St. Aignan’’ in October. Also, a 
cursory glance through articles on insects in Novitates Zoologicae in 


M. LeCroy & W.S. Peckover 230 Bull. B.O.C. 1998 118(4) 


1898 and 1899 (Rothschild & Jordan 1898, Warren 1899) disclosed a 
number of species obtained on “‘St. Aignan”’ in September, October 
and November. 

Meek’s note, added to letter no. 59 and dated 29 November, was 
written from Samarai, his home base and the port from which he would 
have shipped specimens. It seems logical to assume, then, that he had 
left his collectors on Kimuta a few days prior to this and returned home 
to ship the specimens and prepare for his month of cruising about, 
hiring helpers and buying food for his projected stay on Rossel I. 
(letters nos 60 and 63). Labels on specimens dated 26 November-— 
28 December alone are not in Meek’s hand, so we believe that all 
specimens from this period were taken by his collectors on Kimuta. 

In Table 1, we have included in the Misima list species collected by 
Meek from 2 August—4 September; in the Kimuta list species collected 
26 November—28 December. We have listed the specimens collected 31 
July and 5-11 September separately. Analysed in this way, the number 
of species collected by Meek on Misima is 40; Hamlin added four, we 
added ten, Tristram added three, Finch added two, and ‘Tolhurst 
added one, for a total of 60 species known from Misima. Meek collected 
29 species on Kimuta, DeVis added two, for a total of 31 species known 
from Kimuta. 

‘There were 14 species collected a century ago but not observed 
by us. Three, Himantopus himantopus, Alcedo atthis, and Gerygone 
magnirostris, are found near fresh water, of which we saw little. Three, 
Micropsitta pusio, Halcyon saurophaga, and Monarcha guttula, are 
inconspicuous and/or local. Five, Chrysococcyx lucidus, Halcyon 
macleayi, H. sancta, Merops ornatus, and Coracina novaehollandiae, are 
southern migrants and our October visit was late for them. 

Coracina tenuirostris tagulana is considered to be a resident race on 
Misima and Sudest. However, named populations of this species from 
New Guinea’s satellite island should be reassessed, as Mees (1982) has 
synonymized the New Guinea race muelleri with the Australian race 
tenutrostris and considers most mainland New Guinea records, except 
those from the south of the island, to be migrants from Australia. He 
suggests that careful re-examination of specimens from the New 
Guinea islands may show them to be migrants also. 

Meek collected one specimen of Monarcha trivirgatus melanopterus 
on 27 August on Misima and at least 13 on Kimuta, as well as the nest 
and egg Hartert (1899: 208) reported. This subspecies, type locality 
Round Island, a small island off Sudest, is probably also largely a small 
island specialist. Hamlin did not collect it on Misima but found it on 
East and Hastings. 

Ptilinopus rivoli has perhaps actually become rarer due to hunting 
pressure. Meek collected at least six specimens; Hamlin collected one 
during a much shorter stay. Thus, with this possible exception and 
contrary to our first impressions, it is not apparent that any species 
have disappeared from Misima in the last hundred years despite 
massive habitat destruction. But a cautionary note should be sounded, 
for we have no reliable information on changes in abundance of any 
species; and forest species may already be at risk. - 


M. LeCroy & W.S. Peckover 231 Bull. B.O.C. 1998 118(4) 


Taxonomic remarks 


Seven subspecies’ descriptions have been based on Meek’s “St. 
Aignan”’ collection and have Misima as the type locality. These need to 
be re-examined in the light of the above re-analysis. 

Accipiter novaehollandiae misulae. ‘This subspecies was described by 
Mayr (1940: 11). Meek collected four females (including the holotype, 
AMNH no. 832853, Meek no. 858), two males in adult plumage and 
one immature male, during 18-29 August on Misima, which remains 
the type locality. No Meek specimens were found in other museums. 

Collocalia esculenta misimae. Salomonsen (1983: 46) chose as his 
holotype AMNH no. 634526 (Meek no. 789), collected on 11 August. 
Three additional specimens were collected on 1 September, and 
Misima is the correct type locality. One specimen in the BMNH was 
collected on 4 December on Kimuta. 

Alcedo atthis hispidoides. Stresemann (1913: 316) designated AMNH 
no. 636477 as holotype of a now-synonymized form, Alcedo ispida 
pelagica, a specimen collected on 3 September. This holotype is Meek 
no. 953; this number had been incorrectly overwritten as 963, but 953 
is correct, judging by the collecting date. Five other specimens at 
AMNH and one at ROM were collected on Misima 2—30 August. None 
was collected elsewhere and Misima remains the type locality. 

Gerygone magnirostris onerosa. Hartert (1899: 209) named G. 
rosseliana onerosa based on Meek’s material. The holotype, AMNH 
no. 606581 (Meek no. 964), and another specimen were collected on 
5 September; a third specimen was collected on 9 August. Hamlin 
collected the species on Misima, but it has not been reported from the 
smaller islands. We believe that the 5 September specimens came from 
Misima and do not recommend amending the type locality of this 
subspecies. 

Pachycephala pectoralis collaris. Rothschild and Hartert (1918: 311) 
named P. p. misimae, now a synonym of P. p. collaris. Galbraith (1956: 
203, 206-209) synonymized both rosseliana and misimae with collaris 
and considered Misima birds somewhat intermediate between the other 
populations. Differences are slight within the island populations, with 
Rossel birds more distinct, and Coates (1990: 211) has recognized 
rosseliana. P. p. collaris has been collected on Kimuta and in the 
Deboyne Group by others; according to Coates (1990: 211) it is 
confined to small islands in the Marshall Bennett, Bonvouloir, Conflict, 
Deboyne and Renard groups and Teste Island. 

The holotype, AMNH no. 657996 (Meek no. 1044), was collected on 
29 November on Kimuta and the type locality should be amended 
accordingly. Six additional AMNH specimens were collected on 
31 July, 26 November, and 3, 4, 7 and 20 December. The ROM has 23 
more Meek specimens from the following dates: 31 July (two), 
10 September (three), November (two), and December (16). The 
BMNH has four more, collected 1, 17, and 28 December. These are 
from Kimuta and the small island where Meek collected on 31 July. 
The Fleming specimens in the ROM and AMNH are all marked 
“cotype’ (=syntype), but Rothschild and Hartert designated a 


M. LeCroy & W.S. Peckover 232 Bull. B.O.C. 1998 118(4) 


holotype. The other specimens in AMNH from the Rothschild 
Collection would be paratypes, but the Fleming and BMNH specimens 
have no nomenclatural standing. ‘The Kimuta specimens are, however, 
topotypes. 

Zosterops griseotinctus griseotinctus. Hartert (1899: 210) named 
Z. aignant (=Z. g. griseotinctus). ‘The lectotype (Hartert, 1920: 436), 
AMNH no. 700680 (Meek no. 1132), was collected by Meek on 
7 December, and the type locality should be amended to Kimuta. 
Seven additional AMNH specimens (paralectotypes) were collected on 
31 July, 5 September (four), 30 November and 17 December. Four 
ROM specimens and four BMNH specimens, all collected in 
December, were not seen by Hartert and have no nomenclatural 
standing. They are topotypes, as they are all from Kimuta. DeVis also 
reports this species from Kimuta and Hamlin from the Bonvouloir, 
Deboyne and Conflict groups. It has not been reported from Misima. 

Mees (1961: 131-143) discusses the Zosterops griseotinctus group in 
some detail and synonymizes Z. g. aignani with Z. g. griseotinctus. He 
recognizes, as a poorly defined subspecies, Z. g. longirostris, including 
the specimens collected by Hamlin in the Bonvouloir Islands. 
Subspecies limits, and Mees’ discussion (on p. 135) of the source of the 
similar but widespread small island populations that he includes in 
the species Z. griseotinctus, need to be reconsidered in the light of the 
discovery that the species does not occur on Misima itself. Perhaps they 
came, as Mayr (1944: 169, 1955: 44-45) has suggested, “‘ultimately 
from ‘Torres Strait’; or even by westward expansion of some of the 
Solomons’ species included by Mees in the griseotinctus group. 

Myzomela albigula pallidior. Myzomela pallidior was named by 
Hartert (1989a: 21). In the original description, wing measurements 
of a male and a female are given. Hartert (1919: 174) designated the 
male as the lectotype, AMNH no. 692674 (Meek no. 725), collected on 
31 July. According to our analysis, the type locality is not Misima, but 
it is uncertain which of the small islands between Basilaki and Misima 
it came from. We are not aware of any collections having been made in 
the Engineer Group, so do not know whether or not the species occurs 
there. There is only one female in AMNH, which becomes the 
paralectotype; it has Meek’s no. 1178, and was collected 13 December 
on Kimuta. Four other males, all collected 4-17 December, are from 
Kimuta. The BMNH has two specimens collected 9 and 17 December. 
Hamlin collected it on small islands in the Bonvouloir Islands and in 
the Deboyne Group, and it has not been found on Misima. 


Remarks on other species 


In addition to those discussed below, we netted, photographed, and 
released the following species: Accipiter novaehollandiae, Accipiter 
poliocephalus, Lorius hypotnochrous, Rhipidura rufiventris, Monarcha 
guttula, Myiagra alecto, Dicaeum pectorale, Nectarinia aspasia, 
Myzomela mgrita, and Aplonis metallica. 

Egretta intermedia. We saw a medium-sized egret with yellow bill 
and black legs and feet feeding in a marshy area near Bwagaoia. ‘This is 
a first record for Misima. 


M. LeCroy & W.S. Peckover 233 Bull. B.O.C. 1998 118(4) 


Haliastur indus. Even though Richards collected a specimen, now in 
LIVCM, on 12 October 1879, it was not listed by Tristram (1882). 
Hamlin’s specimen, collected on Misima on 23 July 1930, also remains 
unpublished. These two specimen records, along with our sight record, 
constitute the first published record of the species for Misima. 

Haliaeetus leucogaster. We were first told of the occurrence of this 
species by A. Stevens and later saw it at several locations. 

Falco peregrinus. Alan Stevens reported having seen a pair. 

Megapodius freycinet. We were shown a mound, opened and emptied 
of eggs, in a recently cut forest area, with the debris still being burned. 

Pluvialis squatarola. Meek collected three specimens of Pluvialis 
fulva on 10 September and 18 December, probably all on Kimuta. A 
fourth Pluvialis, collected by Meek on 20 December on Kimuta and 
published by Hartert as P. fulva, is actually a specimen of P. squatarola, 
showing the characteristic white rump and black axillaries of that 
species. That most of Meek’s shorebird specimens were from Kimuta 
reflects both that Misima has few beaches and that the timing of his 
stay on Kimuta was better for northern migrants. 

Sterna sumatrana. Meek collected a specimen on 23 December on 
Kimuta that was overlooked by Hartert. Hamlin reported ‘‘sumatras 
and noddies’’ common about Misima and in the Conflict and Deboyne 
groups. 

Anous sp. As noted above, Hamlin reported noddies. 

Chalcophaps indica. Hartert (1899: 214) declared this species to 
be very common. However, Meek’s six specimens were collected on 
9 September and in November and December, perhaps all on Kimuta. 
Hamlin did not collect it. We had one brief glance at an individual as it 
flew between patches of vegetation near Bwagaoia. Its rarity in the areas 
we visited may be due to the high human population density, as this is 
a desirable food species. 

Caloenas nicobarica nicobarica. Meek noted that the iris was 
yellowish—white in all three of his specimens. This differs from the grey 
iris usually noted for this species. Photographs of the species made on 
Tench Island by WSP show some individuals with a grey iris and some 
with a yellowish iris. In the AMNH, specimens having soft parts noted 
are about equally divided between a whitish-yellow and a grey or 
grey—brown iris, variable throughout the range and not related to sex of 
the bird. Young birds tend to have a brown iris. Other colours recorded 
were red or orange, umber, and two-tone with the outer ring grey and 
the inner tan (once). 

Ducula pinon salvadori. We saw this species and frequently heard its 
low-pitched calls. A young bird with feathers in sheath was brought in 
to us and photographed. Its plumage was identical to that of the adult; 
but the facial skin was bluish, unlike the red of adults. 

Tristram (1882: 996) named this form Carpophaga salvadori from 
specimens collected on Misima by G. E. Richards. One of these, a 
male, is now in LIVCM, no. T.10054, collected on 9 October 1879. 
Wagstaffe (1978: 8) noted that ‘““Tristram, when describing this bird, 
had before him three specimens, one male and two females, but only 
the above specimen is listed in his catalogue and as a type. There are 


M. LeCroy & W.S. Peckover 234 Bull. B.O.C. 1998 118(4) 


two specimens in the British Museum, collected by Richards from the 
same locality and these are possibly the other syntypes.’’ However, 
neither of the latter is listed by Warren (1966). 

Robert Prys-Jones (im litt.) informs us that there are two specimens 
in BMNH that are most probably the ones referred to above. 
No. 1889.2.12.129 was “‘acquired via the Salvin/Godman collection. In 
addition to their label, it carries what I presume to be the original label 
(written on paper from “The Australian’!) stating: St. Aignan, 
Louisiade Archipelago, Capt. Richards R.N. No sex/age is given, but 
the register states ad.’ No. 188.11.14.20 ‘‘only carries a B.M. label 
which says merely Louisiade Archipelago, ex ‘Tristram. No mention of 
a collector. However, the register shows it to be part of a batch 
purchased off Tristram and collected by ‘Lieut. Richards’. Old string 
on the legs of the specimen suggests a label may have been lost.”’ 
Listing of the male as the type by Tristram (1899c: 43) in his published 
catalogue serves to designate it as the lectotype and implies that the two 
BMNH specimens are paralectotypes [International Code of Zoological 
Nomenclature, Article 74(a)]. 

[Ducula zoeae. Hartert (1899: 213), without comment, lists this 
species as having been collected on Misima on 25 July. No specimen of 
the Zoe Imperial Pigeon from Misima or any other island in the 
Louisiade Archipelago is now in the AMNH, nor was one catalogued 
with the Rothschild Collection. Rothschild and Hartert (1901: 113) do 
not list a Misima specimen but do report a specimen (now in AMNH) — 
collected by Meek on Basilaki Island on 25 July 1897. We believe that 
Hartert (1899) inadvertently included the Basilaki specimen and that 
this species should be removed from the Misima list. ] 

Trichoglossus haematodus micropteryx. Meek’s specimens, now in 
AMNH, ROM and BMNH, were collected on 8 September (five), 
29 November, 4 and 11 December, most likely all on Kimuta. Neither 
Hamlin nor we encountered it, and it is difficult to believe we could 
have missed this noisy species had it been present on Misima. We 
believe it is confined to small islands in this area. There is some 
evidence of small island populations differing subspecifically from 
adjacent mainland populations (Diamond & Lecroy 1979: 509) and 
exact information on collecting localities and weights might help clarify 
this distribution puzzle. 

Lorius hypoinochrous hypoinochrous. This species does not appear in 
Hartert’s (1899) annotated list of Meek’s collection, and in fact he says 
“not sent by Meek” in his introductory remarks, but it does appear on 
his appended list of species known from Misima (1899: 216), based on 
Tristram (1882) and DeVis (1890). This surprised us very much, as it is 
perhaps the commonest species on the island. However, there are three 
males and three females in AMNH collected by Meek on Misima 
between 12 and 23 August, and Hartert (1898b: 530) discussed the 
Misima specimens in his paper on Meek’s collection from Sudest 
Island. 

Hamlin collected a male on 25 July; we heard and saw these birds in 
large numbers wherever we were on Misima. In the vicinity of Gulewa 
flowering coconut palms appeared to be a major food source for them, 


M. LeCroy & W.S. Peckover 235 Bull. B.O.C. 1998 118(4) 


and it is possible that they are now more common than previously 
due to the establishment and expansion of coconut plantations in the 
first half of the 20th century. We made tapes of their unusual cat-like 
calls. 

Cacatua galerita triton. ‘This very common species is a great 
agricultural pest, invading the gardens in large numbers. In one case, 
we counted a flock of 20 as they took off from a garden area. ‘They dig 
up manioc tubers, and people are forced to cover young pineapples 
with plastic bags to try to deter this species. Other avian pests in the 
gardens were Porphyrio porphyrio and Corvus orru. 

Scythrops novaehollandiae. We saw one individual flying over Gulewa 
on 5 October, presumably on its southward migration. This is a first 
record for Misima. 

Myiagra alecto lucidus. We saw a female clinging to the trunk of a 
coconut tree, feeding. 

Nectarinia aspasia christianae. Tristram (1889b) described this taxon 
(as Cinnyris christianae) based on two males collected by ‘Thomson on 
Misima. One syntype, 1.15504, collected on 20 October 1888, is now 
in LIVCM (Wagstaffe 1978: 22). The other syntype, formerly in the 
York Museum, is now in the BMNH, no. 1945.53.8, and was also 
collected on 20 October 1888 (Warren & Harrison 1971: 108). It is a 
very common species on Misima in all habitats; Meek also collected it 
on 10 September, and Hamlin reported seeing it in the Conflict and 
Deboyne groups. We saw one partial albino with scattered white 
feathers on the head and three all white tail feathers. 

[Myzomela rubro-cucullata. This form was collected by G. E. 
Richards and described by ‘Tristram (1889a: 228) as being from St. 
Aignan; the holotype is in LIVCM, no. ’T.10244. Salomonsen (in 
Paynter 1967: 360) believed that the locality was an error for Samoa 
Islands and equated it with M. c. nigriventris. Wagstafte (1978: 22), 
however, says that the specimen “‘was examined by R. Sims at the 
British Museum in 1958 who confirmed its identity as M. c. cardinalis. 
No original field label is present but if Tristram were correct regarding 
the collector then it would have been collected in the Southern New 
Hebrides which Richards visited, not in Samoa.’ It was certainly not 
from Misima.] 

Aplons cantoroides. Meek collected one male and three females on 
8 September on a “‘small island off St. Aignan’’, probably Kimuta. 
These are the only specimens in the entire collection that have a locality 
other than “St. Aignan”’ on the label. Meek did not obtain this species 
on Muisima, but obtained an additional 11 specimens on Kimuta in 
November and December. We saw it near Bwagaoia, and six 
individuals stayed around some dead trees in the garden above Gulewa, 
calling and flying about as though preparing to nest. 

The following two species were seen by us but not positively 
identified: 

A rail-like bird ran zigzagging along the edge of a rotten fallen tree 
trunk in the forest. It seemed rather crouched, with its head stretched 
out horizontally. Only the reddish-brown head and blackish back were 
seen before it disappeared. Attempts to flush it again were unsuccessful. 


M. LeCroy & W.S. Peckover 236 Bull. B.O.C. 1998 118(4) 


The general size and description point to Rallina tricolor, but the 
species has never been collected in the Louisiades. 

A medium-sized kingfisher was seen perched in bright sunlight on a 
low branch over the river. It was a solid, almost royal, blue with no 
indication of scaling on the head and the underparts were a rich rufous. 
When it flew, the only turquoise visible was in the rump. The bill was 
long, black and sturdy with a conspicuous red strip at the base of the 
lower mandible, and a white marking on the neck was very obvious. It 
resembled Alcedo azurea, which has not been reported from the 
Louisiades, except that azurea has no turquoise in the rump and 
neither the literature nor specimens indicate the presence of red at the 
base of the lower mandible. 


Acknowledgements 


We were able to visit Misima through the courtesy of the Misima Mines Pty. Ltd. and 
Placer Pacific Ltd., especially with the assistance of Ron Hiatt. The following mine 
personnel were extremely helpful to us while we were on Misima, allowing us to use mine 
facilities and personnel transport (‘“‘manhauls’’): Poate Edoni, Ellis Illaia, lan Lewis, Don 
Reid, Allan Storck, Alan Stevens, and Isreal Isreal, as was Len Selly of Poon Catering. 
We are especially grateful to Alan Stevens for his reports of a pair of Peregrine Falcons 
and a White-bellied Sea Eagle on the island, as well as other birds. 

In Gulewa, Artu Sodias was very helpful in providing accommodation and assistance. 
Both he and Raynold Joshua helped us daily in the field and Darcy Siguia was of 
assistance in Gulewa. 

Our thanks also to Beresford and Anne Love of Port Moresby for their ongoing help 
and friendship, and to Navu Kwapena of the Department of Conservation for his 
continued interest and help. 

Michael Quinnell, Anthropology Department, and Heather Janetski, Bird Depart- 
ment, Queensland Museum, tracked down relevant papers and specimens. Mark Peck 
and Brad Millen at the Royal Ontario Museum, Toronto, provided complete information 
on the Fleming Collection. At The Natural History Museum, London, J. C. Thackray, 
Archivist, and his assistants were very helpful during research on the Meek letters and 
Phil Ackery, Department of Entomology, replied to our queries concerning butterfly 
specimens; Robert Prys-Jones and Michael Walters at Tring searched out information on 
the Meek Misima specimens housed in The Natural History Museum, and answered our 
queries concerning the specimens collected by Richards and Thomson. In addition, 
Prys-Jones read drafts of this manuscript and made many very helpful suggestions, and 
Waiters informed us of the published Tristram type list. Clemency Fisher and Anthony 
Parker, Liverpool Museum, kindly provided us with information on Meek, Richards, and 
Thomson specimens in their care; and Chris Milensky found Meek specimens in the 
National Museum of Natural History in Washington. David W. Snow and Chris Feare 
read the manuscript and suggested useful changes. To all of these we are most grateful. 

The Library Staff at AMNH, Donald Clyde and Roscoe Thompson in particular, were 
always helpful, as were Diane Treon and Maria Rios, Ornithology Department, and Dan 
Rutter, Graphics, who prepared the maps. Thanks also to Lauren May and Ramsey 
Togo, whose computer expertise is most appreciated. Brian Coates’ two fine volumes 
(1985, 1990) have greatly facilitated our research. 


References: 

Brass, L. J. 1959. Results of the Archbold Expeditions. No. 79. Summary of the Fifth 
Archbold Expedition to New Guinea (1956-1957). Bull. Amer. Mus. Nat. Hist. 118: 
1-69. 

Coates, B. J. 1985. The Birds of Papua New Guinea, including the Bismarck Archipelago 
and Bougainville. Vol. 1. Non-passerines. Dove Publications Pty. Ltd., Alderley, 
Australia. 

Coates, B. J. 1990. The Birds of Papua New Guinea, including the Bismarck Archipelago 
and Bougainville. Vol. 2. Passerines. Dove Publications Pty. Ltd., Alderley, 
Australia. ‘ 


M. LeCroy & W.S. Peckover 237 Bull. B.O.C. 1998 118(4) 


DeVis,.C. W. 1890. Reports on birds from British New Guinea. Appendix G. In British 
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Diamond, J. M. & LeCroy, M. 1979. Birds of Karkar and Bagabag Islands, New Guinea. 
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Finch, B. S. [Compiler] 1985. Noteworthy observations in Papua New Guinea and 
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Galbraith, I. C. J. 1956. Variation, relationships and evolution in the Pachycephala 
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Addresses: Mary LeCroy, Department of Ornithology, American Museum of Natural 
History, New York, NY 10024, U.S.A. William S. Peckover, 14 Balanda Street, 
Jindalee, Queensland 4074, Australia. 


© British Ornithologists’ Club 1998 


A parsimonious phylogenetic tree for the 
swifts, Apodi, compared with DNA-analysis 
phylogenies 


by Fan Holmgren 
Received 12 February 1997 


In recent years, new and independent data for evaluating phylogenies 
in birds have been produced through biochemical and statistical 
analyses of genetic structures. The techniques used are complex, and 
room for errors of methodology and ambiguities of interpretation 
should be admitted (Sarich et al. 1989, Lanyon 1992). However, to a 
substantial extent the new results are consistent with one another, but 
are sometimes in conflict with traditional views (Bleiweiss et al. 1994, 
Lee et al. 1996). 

The swifts as a group generally seem to exhibit slow, continuous 
adaptations for their aerial life, giving the impressions of a morpho- 
logical conservatism caused by the uniformity of the food niche. The 
great variability in body size may perhaps be linked with interspecific 
competition. A great deal of other morphological variation may 
probably be linked with the intermittent contacts with solid matter in 
breeding and roosting. Here I present a parsimonious phylogenetic tree 


JF. Holmgren 239 Bull. B.O.C. 1998 118(4) 


- Afrotropical 

origin 1236 11, 12 138 Apodini 
Apus 

Characters found Tachymarptis 

in Cypseloidini: 
Cypsiurus 

Two carotid 

arteries Aeronautes, 
Panyptila, Tachornis 

No use of 

salivainnest _ Schoutedenapus 

building 

9 10 Chaeturini 

Broad sternum, Hirundapus 

slender humerus 
Chaetura 

Diastataxic 

wing Mearnsia, Zoonavena,, 
Telacanthura, 

Naked hindlimb, Rhaphidura, Neafrapus. 

anisodacty] foot 

7itts Collocaliini 
Aerodramus 
Collocalia 
Hydrochous 
4 5 Hemiprocnini 
Hemiprocne 
3 Cypseloidini 
: Cypseloides 

Streptoprocne 


Figure 1. Hypothetical phylogenetic tree for the swifts. No time scale. Numbers denote 
change of characters as follows: 1. One carotid artery. 2. Use of saliva in nestbuilding. 3. 
Gradual change towards narrow sternum and stout humerus as adaptations for aerial life. 
4. Nesting and roosting on branches, perching. 5. Near stasis (perhaps even regression) in 
change of sternum and humerus owing to less aerial life. 6. Eutaxic wing. 7. Nesting and 
roosting in caves, some have feathered hindlimbs. 8. Echolocation. 9. Nesting and 
roosting inside hollow trees, square and spined tails. 10. No nestbuilding. 11. Feathered 
hindlimbs. 12. Nesting and roosting in foliage of trees, reduced numbers of phalangi, 
reversed hindtoe, toes in opposite-pairs. 13. Change to more solid nest and roost sites, 
tendency towards “all four toes forwards’’. 


for the swifts, which is consistent with a hypothetical historical 
narrative (Mayr 1988) and zoogeographical facts. This simple tree may 
serve as a null hypothesis: suggested instances of further convergent 
or parallel evolution in the specified characters should be explicitly 
analysed. 


Methods 


The suggested phylogenetic tree in Figure 1 is derived from key 
characters (two carotid arteries or one carotid artery (Glenny 1955, 
Sibley & Ahlquist 1990), no use of saliva or use of saliva in nestbuilding 


J. Holmgren 240 Bull. B.O.C. 1998 118(4) 


(Marin & Stiles 1992, Chantler & Driessens 1995), gradual change 
towards narrow sternum and stout humerus as adaptations for aerial 
life (Sclater 1865, Harrison 1984), diastataxic (i.e. with apparent 
absence of 5th secondary remex) or eutaxic wing (Clark 1906, Sibley & 
Ahlquist 1990)) linked with a general understanding of swift evolution, 
taking into account the role of various behaviours and substrata in 
breeding and roosting. A similar approach has proved successful in the 
swallows (Winkler & Sheldon 1993). The clarification of nest building 
in the Cypseloidine swifts (Marin & Stiles 1992) is an important recent 
achievement. The first methodical survey of all swift species (Chantler 
& Driessens 1995) facilitated an overview. 

In any phylogenetic tree, any lineage from the ancestral species to a 
recent species may be envisaged as a straight continuous line, with all 
other lineages branching off from that arbitrarily chosen trunk of the 
tree. In order to standardise trees for the swifts, I suggest that Apus 
apus should be chosen as this favoured recent species, simply because 
this will produce trees that are most similar to the traditionally accepted 
order between the main groups of swifts, and so will make direct 
comparisons easier. Since only the Cypseloidine swifts show primitive 
states in all the key characters, they are placed at the base of the 
hypothetical phylogenetic tree. 


Results and discussion 
The hypothetical phylogenetic tree—Figure 1 

Brooke’s (1970a) division of the Apodidae into the subfamilies 
Cypseloidinae and Apodinae (instead of the then traditional division 
between the Chaeturinae and the Apodinae, based on differences in the 
hindlimbs, derived from Sclater’s (1865) Chaeturinae and Cypselinae) 
was based on differences in a number of characters. One of Brooke’s 
Cypseloidine characters (diastataxic wing) is also present in the 
treeswifts, which I now suggest should be placed in Apodinae. ‘Two 
characters (anisodactylous feet, downlike semiplumes on young 
nestlings) are present also in some Apodine species. For three 
characters (egg white profiles with four peaks, little or no transpalatine 
processes, simple Musculus splenius capitis) there is uncertainty about 
their exact validity, and one character (that there is a sexual 
dimorphism of plumage in some Cypseloidine species) is questionable 
since it may be an age character (females acquiring adult plumage later, 
M. Marin, pers. comm., Chantler & Driessens 1995). However, 
Brooke’s division into subfamilies still seems warranted by two of his 
characters: the presence of two carotid arteries (Glenny 1955) and the 
now confirmed (Marin & Stiles 1992) lack of saliva in nest building in 
the Cypseloidine swifts. 

Sibley & Ahlquist (1990) critically reviewed morphological charac- 
ters used in phylogenetic work. Generally, conditions of the carotid 
arteries and the 5th secondary were found to be useful to some degree, 
but not consistent. However, within groups, two carotid arteries and 
diastataxy were consistently found to represent the ancestral condition. 
Use of saliva in nest building is special for swifts, and thus represents 
the derived condition. 


F. Holmgren 241 Bull. B.O.C. 1998 118(4) 


The treeswifts have one carotid artery, and they use saliva in nest 
building. Hypothetically, I suggest that the treeswifts should be treated 
as a tribe Hemiprocnini within Apodinae, so all extant swifts belong to 
the family Apodidae. Differences in anatomical details (Lowe 1939) and 
plumage in the treeswifts may well be explained by their different 
lifestyle. Fossil and osteological evidence (Sclater 1865, Harrison 1984) 
show a gradual change of sterna and humeri in the swifts, caused by 
selection forces linked with their extremely specialised aerial life. A 
broad sternum and slender humerus in the treeswifts may be explained 
by near stasis (perhaps even some regression) in this change, following 
the adoption of a less aerial life. 

Analysis of changes in humeral structures led Karkhu (1992) to quite 
different conclusions, suggesting long independent evolution of the 
Hemiproenide and the Apodide swifts. He even suggested that the 
treeswifts should be placed in a suborder Hemiprocni, while the sub- 
order Apodi would include the other swifts and the hummingbirds, 
Trochilidae. He did not consider, however, the possibility of near stasis 
in the treeswifts, coinciding with convergent changes in other lineages 
leading to recent taxa. His interesting analysis of functional causes for 
the observed changes may be interpreted in support for convergent 
change. 

The Cypseloidine and Hemiprocnine swifts have diastataxic wings, 
while all other swifts have eutaxic wings (Clark 1906). 

In Collocaliini the Giant Swiftlet Hydrochous gigas is remarkably 
similar to the Cypseloidine swifts, and its choice of nest site is similar 
(M. Marin, pers. comm., Somadikarta 1968, King 1987). It cannot 
echolocate, and its nest is similar to Cypseloidine nests, except that 
some saliva is used (Becking 1971). 

When erecting the genus Schoutedenapus, De Roo (1968) pointed out 
a close similarity to Apus, but he found it impossible to place the taxon 
within Apus owing to the unmodified (anisodactyl) feet. He discussed, 
however, the possibility that the feathering of the legs may have been 
a first step in the evolution of the Apodine foot. Brooke (1970a) 
temporarily placed Schoutedenapus in Collocaliini, and he is followed 
by Chantler & Driessens (1995). In agreement with De Roo, I contend 
that the feathering of the hindlimbs in Schoutedenapus may be a first 
step in the evolution of the Apodine foot. Important evidence is still 
missing; for example no skeleton has been examined (C. T. Collins, 
pers. comm.). As it is, I suggest that MSchoutedenapus should 
temporarily be included, as an early diverged taxon, in Apodini. 


The historical narrative 

If the birds, as seems likely (Futuyma 1986, Chiappe 1995; for a good 
overview, though discordant in conclusions, see Feduccia 1996), 
evolved from a bipedal running and jumping insectivore catching prey 
in the air with the mouth (Caple et al. 1983), the swifts in principle 
might be the living representatives of a continuous trajectory, 
increasingly refining the faculty of flight, all the time using similar 
food: flying insects. DNA analyses suggest that the hummingbirds 
diverged from the swifts about 95 million years ago (Sibley & Ahlquist 


3}. Holmgren 242 Bull. B.O.C. 1998 118(4) 


1990), obviously changing to a different food niche. The earliest 
records in a revision of fossil swifts (Harrison 1984, see also Feduccia 
1996) are from Europe and about 50 million years old. 

The swallows have had a shorter time than the swifts for the 
adaptations to their aerial life. According to DNA analyses (Sibley & 
Ahlquist 1990) the passerines diverged about 100 million years ago, 
with the swallows diverging after about half that time. Both swallows 
and swifts seem to have evolved in and radiated from the Afrotropical 
region (Brooke 1970b, Turner & Rose 1989). One might have expected 
a more complex pattern for the swifts, considering their more ancient 
origin. Perhaps earlier radiations of swifts were largely wiped out in the 
great extinction about 65 million years ago (not unrealistic, if it was 
caused by an extraterrestrial impact, Alvarez et al. 1980), while one or 
a few species survived in the Afrotropics, and so had a chance to 
become ancestral to all later forms. 

Winkler & Sheldon (1993) superimposed the nest-building behav- 
iours on a DNA-hybridization phylogeny for 17 species of swallows. 
A remarkable evolutionary conservatism was revealed. Three main 
groups are characterised by distinct methods of nest construction. Each 
method seems to have evolved just once, and the three groups have 
largely distinct geographical distributions. Those that excavate burrows 
for breeding are in the Afrotropics, except Cheramoeca in Australia, and 
Riparia, which has spread to the northern continents. Those that 
typically adopt cavities are in the New World, mainly tropical America, 
except Psalidoprocne fuliginosa and Phedina borbonica in the Afro- 
tropics. In those that typically build mud nests, Hirundo species are 
spread almost globally, while Delichon is restricted to Eurasia. The 
Tree Martin Hirundo nigricans in Australia may seem a cavity adopter, 
but uses mud on occasions, sometimes building a full mud nest. In 
contrast to earlier hypotheses (Mayr & Bond 1943, Turner & Rose 
1989), burrow excavating was found to be a more primitive state than 
cavity adoption. 

A similar evolutionary conservatism is evident in the five main 
groups of swifts, which use five distinct substrata for breeding and 
roosting: (1) the ground, (2) branches of trees, (3) caves, (4) the 
inside of hollow trees, (5) foliage of trees, holes in trees, crevices in 
cliffs, etc. Similarly, the five groups have largely distinct geographical 
distributions. 

The first group, the Cypseloidinae, are restricted to the New World, 
but fossil finds in Europe indicate that they represent an early westward 
radiation from the Old World. They have an ancient method of nest 
building, “‘rooting’’ living vegetable matter, and so are restricted to 
humid sites with some daylight near waterfalls (Marin & Stiles 1992, 
Marin 1997). 

In the other groups, probably uniquely in birds, the use of saliva in 
nest building gave new evolutionary options: to place nests in various 
dry and/or dark sites. 

The treeswifts, in using “‘the outside’’ of trees, may seem to have a 
behaviour with some similarity to that of the Apodine swifts. However, 
several characters indicate that they diverged much earlier than the 


¥. Holmgren 243 Bull. B.O.C. 1998 118(4) 


Apodine swifts, even earlier than Hydrochous, using branches and not 
foliage or holes. Their present distribution is in India and eastwards to 
the Solomon Islands. 

In the Collocaliini, Hydrochous gigas is similar to the Cypseloidine 
swifts, nests in daylight near waterfalls (and accordingly has no 
capability of echolocation), and uses much the same vegetable matter 
as the Cypseloidine swifts for nest building, but with some saliva 
mixed in (Somadikarta 1968, Becking 1971). A few Cypseloidine 
species sometimes nest in dark caves, where they place the eggs on 
ledges without building a nest (Whitacre 1989). The presence of the 
tendency towards using caves in some Cypseloidine swifts may 
strengthen the idea that Hydrochous represents an early eastward 
radiation by Cypseloidine-like swifts, however in an early stage of using 
saliva in nest building, in the course of time becoming ancestral to the 
swiftlets. Cypseloidine-like features can be found in some other 
swiftlets. The Glossy Swiftlet Collocalia esculenta cannot echolocate, 
nests in fairly well lit caves, and uses much vegetable material in its 
nests (Francis 1987, Chantler & Driessens 1995). ‘Tarburton (pers. 
comm.) in Western Samoa observed a small colony of the White- 
rumped Swiftlet Aerodramus spodiopygius nesting in the spray of a 
waterfall in a very light situation on a cliff. Feathering on hindlimbs is 
present in some Collocaliine species and subspecies (Chantler & 
Driessens 1995). 

The Chaeturine swifts, the spinetails and needletails, all typically 
use the inside of hollow trees or similar structures, such as chimneys. 
Their use of the tail as a prop is generally accepted as an explanation 
for their stiff and square tails with spines. Several genera are present 
in the Afrotropics; a few of their species are also found far eastwards. 
The genera Hirundapus and Mearnsia clearly represent eastward 
radiations, while Chaetura, in the New World, represents a westward 
radiation. 

Most Apodine species are known to use solid sites, like crevices and 
holes in cliffs or trees, for nesting and roosting. However, I suggest that 
the original causes for the changes in their hindlimbs (feathering, 
reduced numbers of phalangi in toes III and IV from 4 and 5 
respectively to 3, reorientation of toe I (hindtoe) inwards-forwards, the 
toes forming a pincer-like grasp, toes I and II opposing toes III and 
IV, Collins 1983) are linked with use of foliage of trees for nesting 
(palm swifts, Bock & Miller 1959) and roosting (a behaviour still 
present, for example, in the Common Swift, Holmgren 1993), and not 
primarily with clinging to vertical rough surfaces, as has often been 
claimed. Schoutedenapus (with feathered hindlimbs) and the palm 
swifts (Aeronautes, Panyptila, Tachornis, Cypsiurus, with the toes more 
clearly oriented in opposed pairs than in Apus and Tachymarptis, which 
tend towards “all four toes forwards’’) then may be thought to 
represent early stages in these changes. I suggest that the Apodine 
swifts should be characterised by a gradual change in the hindlimbs, in 
the form of a transformation series, rather than by just the most 
advanced stages of that change, a redefinition that opens up the 
possibility to include Schoutedenapus. They are present in the 


J. Holmgren 244 Bull. B.O.C. 1998 118(4) 


Afrotropics and the Palearctic, but some species spread eastwards, 
and others westwards to tropical America (Aeronautes, Panyptila, 
Tachornis). 

The hypothetical phylogenetic tree in Figure 1, following this 
historical narrative, gives a parsimonious solution, and it suggests few 
changes in the current taxonomy of the swifts (cf. Brooke 1970a, 
Chantler & Driessens 1995). At present, this only means that it is a 
simple model for the understanding of swift evolution, and it cannot 
claim to be faultless. 

In the phylogenetic tree there are two cases of convergent evolution. 
The first one is caused by my insistence on placing the Hemiprocnine 
swifts together with all other swifts that have one carotid artery and use 
saliva in nest building. This forces me to admit gradual change towards 
a narrow sternum and stout humerus in two different lineages. 
However, that tendency is generally present in the swifts owing to 
their aerial life style, so this convergence should be expected to be 
omnipresent in the group, except in the treeswifts. 

‘The second case is the feathering on the legs in some swiftlets, which 
may be convergent with the earliest stage in the evolution of the 
Apodine foot. 


Comparisons with DNA-analysis phylogenies 

In Figure 2 the phylogeny for the swifts presented by Sibley & 
Ahlquist (1990) is adapted to the earlier proposed standard, so it can be 
directly compared with the tree in Figure 1. Agreement with Sibley & 
Ahlquist’s results, with Hemiprocne in a separate family being the 
earliest branching, was found in a repeated study using DNA-DNA 
hybridisation (Bleiweiss et al. 1994). The trees in Figures 3 and 4 are 
adapted from Leeet al. (1996), who used DNA-sequencing. They 
derived the first tree by maximum likelihood estimate based on 
cytochrome b sequences, the second is a bootstrap consensus tree for 
1,000 replicates. Both trees have Hemiprocne in the same position 
as Figure 2. Nevertheless I suggest that this question should be 
regarded as unresolved, until the new molecular techniques are better 
understood. 

Whether the Cypseloidinae are monophyletic seems to be a question 
that requires more precision than is attained at present (different results 
appear in Figures 3 and 4). 

The next problem in Figure 2 is the place of Tachornis. Convergent 
change in the hindlimbs in different lineages is certainly possible 
(cf. the feathering on the legs in some swiftlets). Aeronautes, Panyptila 
and Tachornis might form a separate tribe, representing an earlier 
branching than the Apodini. However, that the (reduced) numbers of 
phalangi in the toes are exactly the same seems to be such a specific 
character that it strongly suggests monophyly. 

Also worrying is the place of Tachornis between the needletails 
and the spinetails. It is possible, but seems unlikely. The place of 
Collocalia nearest to Apus then seems very unlikely, since Hydrochous 
so obviously has several characters in common with the Cypseloidine 
swifts. 


F. Holmgren 245 Bull. B.O.C. 1998 118(4) 


20 


_ 
oO 


Million years ago 


5 
> 
o 
J) 
o 


126! 11'12 18 


(os) 


Apus affinis 


ser ere ee g----- © 


7 0 


Collocalia 


Chaetura pelagica 


Rhaphidura 


Se ee 


172 
Tachornis 


Hirundapus 


Cypseloides 


Streptoprocne 


bo 
i 
o 


Hemiprocne 


ee ee ee 


sebhe wee wee oh eo eww ww ooh @ ow ww ow om 


Figure 2. Phylogenetic tree for the swifts, adapted from Sibley & Ahlquist 1990 figure 
361, derived by DNA-DNA hybridisation. A T,,)H 4.5 million years. Numbers have 
same meanings as in Figure 1. 


Figures 3 and 4 also have Hydrochous near Apus. This would force us 
to infer a very remarkable regression in Hydrochous: loss of using caves, 
loss of echolocation (since it is placed with Aerodramus), minimising the 
use of saliva in nest building, and a renewed use of a behaviour similar 
to that of the Cypseloidine swifts, nesting near waterfalls and using 
similar vegetable matter. 

A remarkable result in Figures 3 and 4 is the polyphyly of the 
Collocaliine swifts. In Figure 3, allowing for only a little uncertainty in 
the applied technique, an arrangement similar to Figure 1, lessening 
the problem with Hydrochous, would appear by connecting the 
Collocalia and the Aerodramus branches, leaving Chaetura on its own. 
This also would fit the ectoparasite evidence mentioned by Lee et al. 
(1996). In Figure 4 the situation is far from clear, and seems to reflect 
difficulties with the applied technique. 


JF. Holmgren 246 Bull. B.O.C. 1998 118(4) 


3 126 1112 18 
: Apus apus 
138 
Apus nipalensis 
Cypsiurus balasiensis 
9 
Chaetura vauxi 
7 8 


Aerodramus spodtiopygius assimilis-V 


loss of using caves, loss of echolocation 
use of ground behind waterfalls 


Hydrochous gigas 
Aerodramus terraereginae 
Aerodramus brevirostris 

- Aerodramus maximus 


Aerodramus sawtelli 
Aerodramus bartschi 


Aerodramus spodiopygius spodiopygius 


Aerodramus elaphrus 
Aerodramus francicus 
Aerodramus fuciphagus germani 
Aerodramus salanganus G 
Aerodramus fuciphagus vestitus* 


Collocalia troglodytes 
Collocalia linchi 

Collocalia esculenta cyanoptila 
Collocalia esculenta marginata 
Collocalia esculenta bagobo 


Cypseloides niger 


Streptoprocne zonaris 
1245 
Hemiproene mystacea 


Figure 3. Phylogenetic tree for the swifts, adapted from Lee et al. (1996) figure 2, 
derived by maximum likelihood estimate based on cytochrome b sequences. No time 
scale. Numbers have same meanings as in Figure 1. 


Conclusions 

Several studies (Prum 1990, Clayton & Harvey 1993, de Queiroz & 
Wimberger 1993, Winkler & Sheldon 1993, Paterson et al. 1995, 
Kennedy et al. 1996) confirm the opinion that behaviour may be as 
reliable as other characters for evaluating phylogenies. In the swiftlets, 
however, Lee et al. (1996) did not find nest characters to be 
phylogenetically reliable. I suggest that behavioural characters in the 
swifts, at a higher taxonomic level, may reveal an evolutionary 
conservatism rather similar to that found in the swallows (Winkler & 
Sheldon 1993). 

DNA-hybridization and other techniques for genetic analyses have 
produced interesting new and independent data for evaluating 
phylogenies. In the swifts more work is required before consensus may 


}. Holmgren 247 Bull. B.O.C. 1998 118(4) 


3-1:2.6 1112138 
--4-}-}----- Apus apus 
78 
Aerodramus spodiopygius assimilis V 
loss of using caves, loss of echolocation 
78 use of ground behind waterfalls 
_ Hydrochous gigas 

Aerodramus terraereginae 
Aerodramus brevirostris 
Aerodramus maximus 


Aerodramus spodiopygius spodiopygius 
Aerodramus sawtelli 

: Aerodramus bartecht 

: Aerodramus salanganus B 

; Aerodramus spodiopygius assimilis S 
: Aerodramus fuciphagus vestitus 
Aerodramus elaphrus 

Aerodramus francicus 

Aerodramus fuciphagus germani 
Aerodramus salanganus G 
Aerodramus fuciphagus vestitus* 


4 Jose po Sessler ee ee Chaetura vauxt 


TO eb ee ee ee ree enn 


fe Pee eee ee Apus nipalensis 
oe ee oo ae Cypsiurus balasiensis 


~] 


Collocalia troglodytes 
Collocalia linchi 

Collocalia esculenta cyanoptila 
Collocalia esculenta marginata 
Collocalia esculenta bagobo 


Cypseloides niger 


Streptoprocne zonaris 
12465 


Hemiprocne mystacea 


’ Figure 4. Phylogenetic tree for the swifts, adapted from Lee et al. (1996) figure 3, a 
bootstrap consensus tree for 1000 replicates. No time scale. Numbers have same 
meanings as in Figure 1. 


be established. The aim should be full analyses of all taxa with all 
available techniques. 


Summary 


A parsimonious phylogenetic tree for the swifts, which is consistent with a hypothetical 
historical narrative and zoogeographical facts, is presented. The treeswifts are treated as 
a tribe Hemiprocnini within Apodinae. Inclusion of the genus Schoutedenapus in Apodini 
is suggested. Three trees based on DNA analyses are adapted in order to facilitate direct 
comparisons. Differences between the four trees are discussed. Further detailed studies of 
the swifts with different available techniques are needed. 


Acknowledgements 


C. T. Collins, M. Marin, D. W. Snow and M. K. Tarburton criticised drafts and gave a 
generous share of knowledge, for which I am most grateful. Special thanks to D. Griffin 
for enduring help and support. 


J. Holmgren 248 Bull. B.O.C. 1998 118(4) 


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Brooke, R. K. 1970a. Taxonomic and evolutionary notes on the subfamilies, tribes, 
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Brooke, R. K. 1970b. Zoogeography of the swifts. Ostrich Sup. 8: 47-54. 

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A.-A. Weller 249 Bull. B.O.C. 1998 118(4) 


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Address: J. Holmgren, Rédhakevagen 23, S-274 33 Skurup, Sweden. 


© British Ornithologists’ Club 1998 


On types of trochilids in the Natural History 
Museum, Tring. |. Amazilia Sumichrasti 
Salvin, in relation to morphology and 
biogeography within the A. beryllina complex 


by André-A. Weller 
Received 7 Fuly 1997 


Within the genus Amazilia, about 30 species (e.g. Peters 1945: 29, 
Walters 1980: 33, Sibley & Monroe 1990: 30) and numerous subspecies 
are recognized. Peters (1945; based on Simon 1921) gives the most 
complete list comprising 81 taxa and further types of uncertain status, 
many of which must be judged from an historical point of view. In 
certain cases only single specimens were available to the describer, and 
some of these were believed to be aberrant individuals or hybrids. 
Moreover, taxonomic work on Amazilia has been mainly restricted 
to selected taxa or species groups (subgenera). In view of their 
heterogeneous treatment in publications before and after Peters’ 
check-list, further examination as presented in this study can throw 
new light on the validity of critical specimens or taxa. Here, I examine 
morphological variation within the Berylline Hummingbird, Amazilia 
beryllina, and ascribe A. sumichrasti to a distinct subspecies. 


Material and methods 


Geographic variation in coloration and morphometric data of Amazilia 
specimens in the Natural History Museum, Tring (BMNH) collection 


A.-A. Weller 250 Bull. B.O.C. 1998 118(4) 


(including types; see Warren 1966) were compared with those of 
specimens from other museums (see Acknowledgements). The analysis 
of differences in mensural characters between populations was based on 
measurements of the bill (proximal end of nasal operculum to tip), 
wings and tail (rectrices 1, 5). As colours in Amazilia species (sub- 
family: ‘Trochilinae) are mainly iridescent, general descriptions of 
plumage patterns are subjective and comparisons were made with types 
or selected individuals. Other colours mentioned in the text refer to 
Smithe (1975). Subadult birds could be easily recognized mainly by the 
brownish borders of the body feathers (especially neck, rump), or 
brownish coloration of the breast and abdominal parts. Measurements 
of such birds were excluded from the analysis of mensural data. 
Information on sex and localities of the specimens were obtained 
from their labels. However, in some cases the phenotypic characters or 
measurements indicated that a specimen was wrongly sexed. Collecting 
sites were located using various maps (scale: 1:4,000,000, 1:1,800,000) 
and monographs (Friedmann et al. 1950, Binford 1989), respectively. 
In order to obtain reasonable sample sizes, specimens from adjacent 
localities were grouped in “‘pools’’ (method after Vuilleumier 1968). 


Results and discussion 


Biogeography and morphology within the A. beryllina group 

The current distribution of the races of A. beryllina centres on the 
Pacific slopes from Mexico to El Salvador. The species is a relatively 
common inhabitant of oak-rich woodlands, forest edges and shrub 
(Howell & Webb 1995). Morphologically, the most obvious geographic 
variation occurs in the dorsal plumage, including the tail. In the 
western states from Sonora to Guerrero, A. b. viola can be regarded 
the least contrasted taxon of the group. The plumage is dark golden 
green, similar to A. b. beryllina, but with a rufous lower belly and 
abdomen. The upperparts have sometimes a certain greyish tinge 
and show dark violet (172 in Smithe 1975) only in the tail coverts 
and rectrices. Some intergradation in the ventral coloration towards 
the nominate form can be observed in Michoacan specimens that 
possess reduced rufous parts abdominally. A. b. beryllina ranges from 
District Federal eastward to Veracruz (here reaching the Atlantic 
slope), and southward to northeastern Oaxaca, with an altitudinal 
distribution between 600 and 3,000m. Concerning the dorsal 
coloration, more variation is recognizable in nominate beryllina than in 
A. b. viola, with copperish to purplish gloss on the lower back and 
rump. Besides, the rectrices in A. b. beryllina are more contrasting 
copper to rufous. 

Within the southern beryllina forms, two striking colour morphs 
exist in the colour pattern of the tail. A. b. lichtensteini shows in both 
sexes a rather light gloss in the rectrices that can be best described as 
chrome-coloured (see Moore 1950); partially, the lateral parts are 
purplish. In contrast to all other races, this taxon inhabits a very 
limited area in western Chiapas that seems mainly to be restricted to 
the slopes of Cerro Brujo. A. b. devillei is the only member of the 


A.-A. Weller 254 Bull. B.O.C. 1998 118(4) 


species that occurs outside Mexico, in the highlands of southern 
Guatemala, E] Salvador and central Honduras. It represents the richest 
coloured race of the beryllina group. Remarkably, the dorsal plumage is 
more golden to bronze-green than in all other forms. In particular, the 
upper tail-coverts and rectrices vary from copperish to intensive purple 
red. A common feature shared with A. b. lichtensteini is the less 
extended (visible), paler chestnut coloration at the wing base than in 
A. b. beryllina and A. b. viola. 

From the arid valley of Rio Motagua, northeastern Guatemala, 
Carriker & Meyer de Schauensee (1935) described a questionable 
subspecies, A. b. motaguae. No validity was given to this taxon by 
following reviewers (e.g. Peters 1945, Land 1970). My examination of 
the type series revealed that the most important distinguishing 
characters (i.e. brownish tips in rectrices) should be mostly counted on 
juvenile or female characters. As there exists a relatively high amount of 
intraracial variation in A. b. devillei, the validity of A. b. motaguae 
should be rejected. Additionally, the southernmost Mexican population 
of A. beryllina, ranging from the coastal Atlantic slopes to the central 
mountainous areas of Chiapas, has been included in A. b. devillei (i.e. 
Friedmann et al. 1950, Howell & Webb 1995). 


Morphological aspects and type locality of Amazilia Sumichrasti (Salvin 
1891) 

Holotype: BMNH 1887.3.22.1865, Santa Efigénia, Tehuantepec, 
Oaxaca, Mexico. 

In its general appearance, the type resembles a highly coloured 
individual of the southernmost beryllina race devillei. Characteristic 
features are the relatively dark glittering green plumage, especially on 
the head, and below extending to the abdomen, the chestnut bases of 
the inner primaries and secondaries as well as the contrasting tail (see 
Salvin 1891). It is of interest that the colour of the latter shows a certain 
purplish gloss instead of only “‘coppery bronze’’ mentioned in the first 
description. As found in other members of the genus, the chin and 
upper throat feathers lack conspicuous white subterminal bars, 
suggesting the type to be a male. 

The type locality is situated in extreme southeastern Oaxaca (Santa 
Efigénia). Thus, it has been believed by some authors to represent an 
extreme western example of A. b. devillei or an intermediate towards 
the nominate form (see below), respectively, although the collecting site 
is in particular close to the range of A. b. lichtensteinz. 

With the single type at hand, it was not possible to detect either 
significant morphological or morphometric differences in comparison 
to other beryllina races. The later comparison of a series of possible 
A. b. devillei specimens from Chiapas with typical representatives of the 
race from Guatemala (holdings of MLZ), with regard to the “‘unique”’ 
features of the Sumichrasti type, revealed evidence for the subspecific 
distinctness of the southern Mexican population of A. beryllina. 


Taxonomic aspects 
The taxonomic history of A. Sumichrasti is comparable to those of 
other single specimens once described as new taxa. Salvin & Godman 


A.-A. Weller 252 Bull. B.O.C. 1998 118(4) 


(1892) and Boucard (1895) regarded it as a valid species, an opinion 
shared by Ridgway (1911). Salvin (1892) and Hartert (1900) even noted 
some similarity with Amazilia Ocait (Gould 1859), a presumed 
intrageneric hybrid of A. beryllina and A. cyanocephala (Berlioz 1932, 
pers. obs.) from Mexico, although Hartert also mentioned distinguish- 
ing features (crown less shining green, throat lacking conspicuous white 
centre, tail copperish, chestnut-coloured under tail-coverts). Appar- 
ently, Simon (1921) ignored the bird, whereas Peters (1945) considered 
A. Sumichrasti to be an aberrant specimen of A. b. beryllina or an 
intermediate between the nominate race and devillez. Since then, 
intergradation between both subspecies has been proposed as the cause 
for the coloration patterns in southern Mexican birds (Friedmann et al. 
1950), regarding the coastal strip of southeastern Oaxaca and Chiapas 
as a hybridization zone (Moore 1950). 

In the latest review on the birds of Oaxaca (Binford 1989), there is no 
indication of the existence of another subspecies within the beryllina 
complex although the sumichrasti type is mentioned (regarded as an 
intermediate of nominate beryllina and devillei, too). Against this 
background, it is not surprising that Sibley & Monroe (1990) cite 
A. Sumichrasti in synonymy with A. b. devilleiz. At least, Howell & 
Webb (1995) treat the southern forms as a peculiar morphological 
group, being aware of the taxonomic uncertainties. As mentioned 
above, the comparison of the type with specimens from other parts of 
southern Mexico including the races deville1 and lichtensteim strongly 
supports the view that the beryllina population of coastal and interior 
Chiapas can be separated as a distinct taxon, Amazilia beryllina 
sumichrastt. 


Diagnosis 

Generally, more grass-green and less shining golden-green than 
A. beryllina devillei and A. b. lichtensteini, in this character resembling 
the northwestern form viola, but underparts entirely green as in the 
former ones. Crown relatively dull. Similar to A. b. devillet in the 
coloration on the base of the inner primaries and secondaries (132A, 
Brick Red; lacking in female), not as prominent as in the nominate 
form. Rump nearly without the purplish gloss of A. 6. devilleiz. No 
differences exist in the upper tail-coverts in both sexes and the rectrices 
in males. Adult females and immatures in general with metallic chrome 
median rectrices as in A. b. lichtensteint (but often with purplish traces); 
outer rectrices more dark chestnut (32) to purplish. Concerning other 
features, sexual dimorphism is restricted to the whitish subterminal 
bars of the throat feathers which are more marked in females. 

In size, A. b. sumichrasti is similar to devillei and lichtensteznz, 
although the wings are slightly longer than in the latter form (Table 1). 
The duller green plumage is typical of the northern distributed 
subspecies that occur along the Pacific coast (A. b. viola) or the central 
and northeastern parts of Mexico, respectively (A. 6. beryllina). 


Range 
A. b. sumichrasti is found in the coastal parts up to the mountains 
of extreme southeastern Oaxaca, southern and north-central Chiapas 


A.-A. Weller 253 Bull. B.O.C. 1998 118(4) 
, TABLE 1 
Ranges, means and standard deviations in morphometric characters of Amazilia beryllina 
subspecies. For geographical reasons, only data of beryllina devillei specimens from 
Guatemala are tabled 


Bill (mm) Wing (mm) Rectrix 1 (mm) Rectrix 5 (mm) 
A. b. devillei 

ein=15 18.5-—21.5 52.5-56.5 25.0-28.5 30.0—34.0 
‘ POS Ov 1 54.44+1.19 27.3 + 0.99 Si 7 = 120 
Bein A =23 19.5-21.5 51.0-53.0 25.5-28.5 28.5—32.0 
; 20 73359057 1 51-8480 275 26.9+ 0.97 30.4+1.19 

A. b. lichtensteini 
Bic. a= 24 19.0-—21.0 49.0-55.0 25.0—28.5 30.0-34.0 
: : 19.8 + 0.60 53.5 + 1.65 26.9+ 0.99 31.6+0.99 
Bim 18 19.0-—22.5 50.0-53.5 25.5-28.5 28.0—32.5 
; 20.7 + 0.89 52.1 + 1:06 27-1 + 0.72 30.2 + 1.03 

A. b. sumichrasti 
Male: n=24 19.0-22.5 53.5-56.5 25.5—-28.5 31.5-34.0 
5 i 20.5 + 0.83 5522-5 0.77 27.2 £0.79 32.6 + 0.66 
Bae n= 14 20.0—22.0 49.5-54.5 26.0—28.5 29.0-32.5 
; 21.2 + 0.64 52.8 + 1.14 27.5 0282 30:5.4£2008 


(Fig. 1), but not known from Guatemala. Northward, it inhabits an 
area as far as Rancho Santa Efigénia (type locality; c. 250 m, 16°27’N, 
94°14'W) and the Tapanatepec region, but possibly does not range into 
the adjacent mountains of northern Oaxaca where there is one record 
of a female identified by A. Weller as A. b. lichtensteint (USNM, 
no. 467908, La Cova). Residential status of A. b. sumichrasti can be 
assumed for the Sierra Madre de Chiapas and probably the Montanas 
del Norte de Chiapas where it reaches an altitude of at least 1,700 m 
(Yerba Buena, 17°06’N, 92°53’W). So far, the northwest border of the 
range remains unclear, as the nominate form has been believed to 
inhabit the parts of eastern Oaxaca that seem to be at least partially 
unexplored (Binford 1989, p. 147). In northwestern Chiapas, it is 
replaced in mountainous areas by lichtensteini, the last recognized taxon 
in the beryllina group (Moore 1950). Apart from the type locality 
(Cerro Brujo, Ocozocuautla), lichtensteini has been believed to range 
also in the central mountains of Chiapas (Friedmann et al. 1950), but 
records of A. beryllina from this region (listed by the data bank of 
ECOSUR, San Crist6bal de las Casas) probably refer to A. b. 


sumichrasti. 


Conclusions 

Presumably, the current population of A. b. sumichrasti once 
connected the beryllina populations from northwestern Mexico to 
Guatemala. For example, clinal variation can be recognized in the 
upper tail-coverts which are reddish purple in A. b. deville: and A. b. 
sumichrasti, but dark purple in the race viola. Based on morphological 
characters, the new taxon is closely allied to A. b. devillet and, with the 


A.-A. Weller 254 Bull. B.O.C. 1998 118(4) 


Rio Grijalva 


v 


Yuxtla Gutiérrez 


17° 


Ciudad 
Ixtepec 
@ 


CHIAPAS 


Presadela _ 


a, Angostura 


PACIFIC OCEAN be, 


e apachula 


? 


95° 94° 935 92° 


Figure 1. Collecting sites (filled triangles) of Amazilia beryllina sumichrasti in Oaxaca and 
Chiapas, southern Mexico. 


difference of the duller plumage and sexual dimorphism in the tail 
coloration, to A. b. lichtensteim. Intergradation of A. b. devillei with the 
nominate form as supposed by Friedmann et al. (1950) and Binford 
(1989) can be excluded not only for reasons of coloration but also of 
distribution, because there is possibly a separation from the most 
northwestern population of the race lichtensteini in Oaxaca. Additional 
distributional data on these subspecies are necessary to determine their 
range boundaries. I presume that A. b. sumichrasti is restricted to the 
southwestern slopes of the Sierra Madre and the Montanas del Norte 
(as indicated by the MLZ series from Yerba Buena, and adjacent 
beryllina localities in the ECOSUR data bank). With the fragmentary 
data at hand, it is problematic to estimate if the swmichrasti populations 
are connected or separated by the valley of the Rio Grialva/Presa de la 
Angostura in interior Chiapas (Fig. 1), where original woodlands have 
been widely cleared (Howell & Webb 1995). 

Altogether, the appearance of morphological characters in the 
southern beryllina subspecies indicates microevolutionary processes as a 
result of isolation of preliminary taxa groups. As has been suggested for 
numerous Central American taxa, including Amazilia species (Howell 
1993), climatic changes in glacial and interglacial periods which caused, 


A.-A. Weller 255 Bull. B.O.C. 1998 118(4) 


e.g., cyclic extension and forcing back of humid forests, may have 
isolated small population groups of proto-beryllina in drier areas. 
Under more suitable environmental conditions, the range could be 
extended leading to some convergence of morphological features. As a 
result, these geohistoric events probably isolated also A. b. lichtensteim 
and possibly the northern population of A. b. suwmichrasti (barrier effect 
of the Rio Grijalva valley?) from the more coastal populations. In 
comparison to the A. beryllina group, similar patterns of distribution 
and morphology exist in other congeneric species found in southern 
Mexico, A. rutila, A. violiceps and A. viridifrons (pers. obs., Howell 
1993). 


Acknowledgements 


I thank the curators and scientific staff of the following institutions for their support of 
my studies: The Academy of Natural Sciences, Philadelphia (ANSP); American Museum 
of Natural History, New York (AMNH); Natural History Museum, Tring (BMNH); 
Field Museum of Natural History, Chicago (FMNH); Museum of Natural Science, 
Louisiana State University, Baton Rouge (LSUMWNS); Moore Laboratory of Zoology, 
Occidental College, Los Angeles (MLZ); Museum of Comparative Zoology, Harvard 
University, Cambridge (MCZ); United States National Museum, Washington, DC 
(USNM); and Zoologisches Forschungsinstitut und Museum A. Koenig, Bonn (ZFMK). 
Particularly, I am obliged to M. Walters for supervising my research at the BMNH. 

Especially I am grateful to K.-L. Schuchmann (ZFMK), who supervised the work. 
He, D. W. Snow, M. Walters and an unknown referee gave helpful critical comments on 
earlier drafts of the MS. Further logistic help was provided by M. Heindl, I. Heynen, 
M. Leutfeld, C. Stein, and U. Wittmann (all ZFMK). Moreover, A. L. Gardner 
(USNM), and P. L. Enriquez Rocha, El Colegio de la Frontera del Sur (ECOSUR), San 
Cristobal de las Casas, Chiapas, contributed valuable information on distributional data. 
Generously, E. Mayr and the MLZ provided accommodation. 

The work in the museum collections was supported by awards or grants from the 
Yessup McHenry Fund (ANSP), Ernst Mayr Fund (MCZ), Field Museum Grant, a 
Frank Chapman Collection Study Grant (AMNH), Smithsonian Research Opportunities 
Fund (USNM), and the Gesellschaft fir Tropenornithologie (GTO). 


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530-534. 

Binford, L. C. 1989. A Distributional Survey of the Birds of the Mexican State of Oaxaca. 
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Boucard, A. 1895. Genera of humming birds. Humming Bird 4: 107-202. 

Carriker, M. A. & Meyer de Schauensee, R. M. 1935. An annotated list of two collections 
of Guatemalan Birds in the Academy of Natural Sciences of Philadelphia. Proc. 
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Friedmann, H., Griscom, L. & Moore, R. T. 1950. Distributional check-list of the birds 
of Mexico. Part 1. Pacific Coast Avifauna 29: 1-102. 

Hartert, E. 1900. Trochilidae. In A. Reichenow (ed.), Das Tierreich, 9. Lieferung. Aves. 
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Howell, S. N. G. & Webb, S. 1995. A Field Guide to the Birds of Mexico and Northern 
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PA. 

Moore, R. T. 1950. A new race of the species, Amazilia beryllina, from southern Mexico. 
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Salvin, O. 1891. Descriptions of new species of Upupe and Trochili in the collection of 
the British Museum. Ann. Nat. Hist. (6) 7: 374-378. 

Salvin, O. 1892. Catalogue of the Picarie in the Collection of the British Museum. Upupae, 
Trochili. 1-433. London. 

Salvin, O. & Godman, F. D. 1892. Biologia Centrali-americana. Aves, vol. 2. London. 

Sibley, C. G. & Monroe, B. L., Jr. 1990. Distribution and Taxonomy of Birds of the World. 
Yale University Press, New Haven, CT. 

Simon, E. 1921. Histoire naturelle des Trochilides (synopsis and catalogue). Encyclopedia 
Roret, L. Mulo, Paris. 

Smithe, F. B. 1975. Naturalist’s Color Guide. American Museum of Natural History, 
New York. 

Walters, M. 1980. The Complete Birds of the World. David & Charles, Newton Abbot, 
London and North Pomfret. 

Warren, R. L. M. 1966. Type-specimens of Birds in the British Museum (Natural History), 
vol. 1. Non-Passerines. London. 


Address: André-A. Weller, Zoologisches Forschungsinstitut und Museum A. Koenig, 
AG Biologie und Phylogenie tropischer Végel, Adenauerallee 160, 53113 Bonn, 
Germany. 


© British Ornithologists’ Club 1998 


Lectotypification of Charadrius rubricollis 


Gmelin, 1789 


by Storrs L. Olson 
Received 9 May 1998 


The note by McAllan and Christidis (1998) regarding the scientific 
name for the Australian bird now known as the Hooded Plover reflects 
a lack of understanding of the rules and procedures of zoological 
nomenclature. ‘The case involves two competing names Charadrius 
rubricollis Gmelin, 1789, versus Charadrius cucullatus Vieillot, 1818. 
MecAllan and Christidis (1998: 60) proposed to resolve what they 
perceived to be “potential confusion’? concerning these names by 
designating a neotype for the former. } 

A neotype, however, is only to be selected in the course of revisory 
work, and then only under exceptional circumstances involving closely 
similar species for which one or both holotypes may be missing, neither 
of which circumstances apply in this instance. Furthermore, a neotype 
is not to be designated as an end in itself, so that the action of McAllan 
and Christidis, whose note has no other purpose, is automatically 
invalidated by provisions of Article 75 (b & c) of the International 
Code of Zoological Nomenclature (I. C. Z. N. 1985—hereafter “‘the 
Code’’). 

The name Charadruis rubricollis Gmelin (1789: 687), was based 
entirely on “‘Lath. syn. III., 1, p. 212, n. 19” which refers to volume 3, 
Part 1, of John Latham’s General Synopsis of Birds (Latham 1785). 
Here Latham described what he called the ‘‘Red-necked Plover’’, this 
being the source of Gmelin’s name rubricollis, neither name being at all 


S. L. Olson 257 Bull. B.O.C. 1998 118(4) 


appropriate for the Australian Hooded Plover, which has no red in the 
plumage. No one in the history of the nomenclatural discussions of this 
species seems to have remarked on this rather obvious fact. 
Appropriateness has no bearing on the validity of a name but it should 
be taken into account when there are other valid grounds for dispensing 
with a misnomer. Latham stated that his Red-necked Plover “‘Inhabits 
the South Seas. Found in Adventure Bay, Van Diemen’s Land 
[T'asmania]”’. 

Mathews (1913: 130) found Latham’s description to be ‘inapplicable 
to every Australian species, and no previous worker had been able to fix 
it on any extra-limital form’’. Latham made no reference to any speci- 
mens or illustrations, so on internal evidence his species would have to 
be regarded as a nomen dubium that is not available for any known 
species. However, the Code suggests that “if an author, in establishing a 
nominal species—group taxon, does not explicitly state what specimens 
constitute the type series, evidence in addition to published evidence 
may be taken into account”’ (recommendation 72B). As this is only in the 
form of a recommendation, some workers may still consider any name 
based solely on Latham’s description to be indeterminable. 

Mathews (1913) went on to build a strong circumstantial case, 
though it is still only that, for Latham’s description being a composite 
based upon two water-colour drawings made by William Ellis on 
Cook’s Third Voyage, these being in the collections of the British 
Natural History Museum and having previously been discussed by 
Sharpe (1906: 205) and subsequently by Lysaght (1959), who used the 
same system for numbering them. ‘The first of these, plate 63, was 
identified by both Sharpe and Lysaght with the species now known as 
the Red-necked or Northern Phalarope, Lobipes lobatus, based on 
Tringa lobata Linnaeus (1758), Ellis’s original specimen having been 
taken “‘between Asia and America’’. The second drawing, plate 67, was 
identified by Sharpe with “Aegialitus cucullatus (Vieill.)’’, based on a 
specimen from “‘Adventure Bay’’. Lysaght (1959: 333) listed this under 
Charadrius rubricollis, following the terminology of Peters (1934), and 
likewise considered that the species depicted is clearly the Australian 
bird now known as the Hooded Plover or Hooded Dotterel. 

Contrary to McAllan and Christidis, Mathews (1913) gave no reason 
for his not using rubricollis for the species in question, although 
inappropriateness may have been more of a factor than their surmise 
that it was because the name was based on two different species. If 
Mathews’ conclusions are accepted as correct, then the name 
Charadrius rubricollis Gmelin, 1789, is a composite, a circumstance that 
arose many times in the compilations of Linnaeus and Gmelin and that 
in no way invalidates the proposed name. The disposition of a 
composite species is resolved by application of Article 74 of the Code: 
“If a type series contains more than one specimen and a holotype has 
not been designated, any author may designate one of the syntypes as 
the lectotype, by the use of that term or an equivalent expression (e.g., 
‘the type’)’’. No action that constitutes lectotypification of Charadrius 
rubricollis Gmelin occurs in any of the literature bearing on this case as 
cited by McAllan and Christidis (1998), however. 


S. L. Olson 258 Bull. B.O.C. 1998 118(4) 


The syntypes of this name are the specimens depicted in Ellis plates 
63 and 67. ‘The fact that these specimens no longer exist is immaterial, 
contra McAllan and Christidis (1998) who mistakenly cite Article 72c 
(v) in this connection. That article, however, applies only when an 
illustration is designated as a holotype but the actual specimen upon 
which it is based still exists. In the present case, the appropriate rule is 
Article 74c: “‘designation of an illustration or description of a syntype 
as a lectotype is to be treated as designation of the specimen illustrated 
or described; the fact that the specimen cannot be traced does not of 
itself invalidate the designation.’ Thus, the paintings themselves in 
effect become the types and McAllan and Christidis (1998:60) are quite 
wrong in stating that ‘“‘no lectotype can be made.” 

McAllan and Christidis (1998:59) misleadingly considered that 
Oberholser (1919) “‘resurrected”’ rubricollis but that he invoked an 
incorrect argument, stating that because “the name rubicollis [sic] refers 
to more than one taxon it would appear to be a case of instant 
homonymy and is thus not valid.’’ However, homonymy involves two 
names with the same spelling being applied independently to different 
species, which is not a factor here. 

Oberholser’s (1919) actions were extremely muddled to say the least. 
He cited Mathews (1913) as showing that Charadrius rubricollis was 
based on drawings of two different species, although he committed a 
rather serious lapsus in stating that one of these was Steganopus tricolor, 
which is a very different species of phalarope, when he meant Lobipes 
(or Phalaropus) lobatus. He went on to conclude as follows: ““The name, 
therefore, should apply [my emphasis] to the species to which the 
greater or most pertinent part of the description refers, which in this 
case is, of course, Charadrius cucullatus. If, however, we take the view 
that it is erroneously described, neither current usage nor the 
commonly accepted codes of nomenclature allow its rejection because 
of indefinite or even erroneous characters, if the description can be 
positively determined as pertaining to a certain species. Thus, in any 
case, we should call the species ordinarily known as Charadrius 
cucullatus Vieillot by the name Charadrius rubricollis Gmelin.”’ 

It is difficult to know what, if anything, can be made of the second 
sentence of this quotation. The description does not apply to “‘a certain 
species’ it applies to two certain species, and nothing in Oberholser’s 
note refers to a “‘type’’ or anything that could be construed as a type in 
such a manner as to meet modern requirements for lectotypification, 
even though Oberholser’s publication has been the only justification 
cited by previous authors for using the name rubricollis Gmelin. 

The name Charadrius rubricollis may yet be regarded as being of 
dubious application if determined solely on the internal evidence of 
Latham’s original description. Latham’s name Red-necked Plover, as 
well as Gmelin’s Latin derivative of it, rubricollis, is clearly indicative of 
what Latham considered the most salient feature of his species and this 
feature does not occur in the Australian Hooded Plover. Furthermore, 
the perfectly descriptive name cucullatus Vieillot was well established in 
the literature of the Hooded Plover prior to 1919 and was used in 
numerous publications subsequent to that date. For these reasons, and 


F. F. Pacheco & B. M. Whitney 259 Bull. B.O.C. 1998 118(4) 


in order to remove any further contention regarding the name, I 
designate the signed drawing by William Ellis listed as number 63 by 
Sharpe (1906: 205) and by Lysaght (1959: 332), depicting a phalarope 
and having the associated information “W. Ellis ad vivum delint: et 
pinxt: 1778. Between Asia and America.’’, as the lectotype of 
Charadrius rubricollis Gmelin, 1789, which then becomes a junior 
subjective synonym of Tvinga lobata Linnaeus, 1758. Consequently, 
the Australian Hooded Plover should henceforth take the name 
Charadrius cucullatus Vieillot, 1818. 


Acknowledgements 


I thank Robert Prys-Jones and Michael Walters, Tring, for reading and commenting on 
a draft manuscript. 


References: 

Gmelin, J. F. 1789. Systema Naturae. 13th edn. G. E. Beer, Liepzig. 

I. C. Z. N. (International Commission on Zoological Nomenclature). 1985. International 
Code of Zoological Nomenclature. 3rd edn. International Trust for Zoological 
Nomenclature, London. 

Latham, J. 1785. A General Synopsis of Birds. Vol. I1I(1). Leigh & Sotheby, London. 

Linnaeus, C. 1758. Systema naturae. 10th edn. Laurentius Salvius, Stockholm. 

Lysaght, A. 1959. Some Eighteenth Century bird paintings in the library of Sir Joseph 
Banks (1743-1820). Bulletin of the British Museum (Natural History) Historical Series 
1 (6): 253-371. 

Mathews, G. M. 1913. The Birds of Australia. Vol. 3, Part 2. Witherby & Co., London. 

McAllan, I. A. W. & L. Christidis. 1998. Neotype of the Hooded Plover Charadrius 
rubricollis Gmelin, 1789. Bull. Brit. Orn. Cl. 118: 59-60. 

Oberholser, H. C. 1919. The status of Charadrius rubricollis Gmelin. Auk 36: 279. 

Peters, J. L. 1934. Check-list of Birds of the World. Vol. 2. Museum of Comparative 
Zoology, Cambridge, Massachusetts. 

Sharpe, R. B. 1906. Birds. Pp. 79-515 in The History of the Collections Contained in the 
Natural History Departments of the British Museum. Vol. 2. British Museum, 
London. 

Vieillot, J. P. L. 1818. Nouveau Dictionnaire d’Histoire Naturelle, 27: 136. 


Address: S. L. Olson, Department of Vertebrate Zoology, National Museum of Natural 
History, Smithsonian Institution, Washington, D. C. 20560. 


© British Ornithologists’ Club 1998 


Correction of the specific name of 
Long-trained Nightjar 


by Fosé Fernando Pacheco and Bret M. Whitney 
Received 13 December 1997 


The specific name of Long-trained Nightjar Hydropsalis (=Macro- 
psalis) creagra (Bonaparte 1850) requires formal correction under the 
articles of the International Code of Zoological Nomenclature (ICZN 
1985). This spectacular nightjar (curiango-tesourao in Portuguese), 
endemic to the southern Atlantic Forest region, was described by 
Nitsch in 1840, with the name Caprimulgus forcipatus. Ten years later, 


J. F. Pacheco & B. M. Whitney 260 Bull. B.O.C. 1998 118(4) 


Bonaparte (1850) named this species Hydropsalis creagra, a name 
clearly understood as a synonym of forcipatus until, for reasons 
unexplained, it was reintroduced by Peters (1940). 

Although relatively few widely distributed papers were published for 
some twenty years following this, the seminal authors working in Brazil 
(e.g., Olivério Pinto, Helmut Sick, and Augusto Ruschi) published 
numerous works employing the correct name Macropsalis forcipata for 
this species. Indeed, almost no papers mentioning the bird were 
published outside Brazil between the appearance of Peters’s volumes 
and Meyer de Schauensee (1966). However, subsequent to the 
publication of Meyer de Schauensee’s (1966) valuable book (based in 
large part on Peters’s published volumes of his Check-list of the Birds 
of the World), the incorrect name Macropsalis creagra seems to have 
become stabilized in the literature. For example, Sibley and Monroe 
(1990) recognized that forcipata had priority as the specific name for 
Long-trained Nightjar, but then suggested that it “‘qualified’’ as a 
nomen oblitum because it had not been used for more than 50 years. 
Even if the ICZN (1985) admitted judgement of a name as a nomen 
oblitum (which it has not since 1 January 1973), this treatment by 
Sibley and Monroe (1990) overlooked pertinent literature published by 
authors in Brazil between 1945 and 1966 concerning this exclusively 
Brazilian species. In accordance with the Principle of Priority (Chapter 
VI, Article 23; ICZN 1985), the correct name for Long-trained 
Nightjar is Macropsalis forcipata (Nitsch 1840). 

We take this opportunity to cite more prominently than has been 
done previously another paper treating a slightly different nomen- 
clatural anomaly concerning the name Hydropsalis brasiliana (Gmelin 
1789), Scissor-tailed Nightjar (curiango-tesoura in Portuguese). 
Teixeira (1992) showed that this name was based on a description and 
a drawing that did not permit identification of the named form as the 
species generally known today as Hydropsalis brasiliana (in fact, some 
points of the description would seem to obviate this identification). As 
demonstrated by Teixeira (1992), the correct name for Scissor-tailed 
Nightjar is best considered Hydropsalis torquata (Gmelin 1789; 
published one page after the name brasiliana). As with the case of 
Macropsalis forcipata, the name Hydropsalis torquata (reintroduced by 
Schneider [1938] and adopted by Peters [1940]) was used consistently 
in Brazilian literature for the Scissor-tailed Nightjar prior to the 
publication of Meyer de Schauensee (1966). 


References: 

Bonaparte, C. L. J. L. 1850. Conspectus generum avium. Tom. I. Lugduni Batavorum: 
Academiae Typographum. 

I.C.Z.N. [International Code of Zoological Nomenclature] 1985. 3rd Ed. International 
Trust for Zoological Nomenclature, London. 

Meyer de Schauensee, R. 1966. The species of birds of South America and their distribution. 
Academy of Natural Sciences of Philadelphia, Narberth, Pennsylvania. 

Nitsch, C. L. 1840. System der Pterylographie. Eduard Anton, Halle. 

Peters, J. L. 1940. Check-list of the birds of the world. IV. Museum of Comparative 
Zoology, Cambridge, Massachusetts. 

Schneider, A. 1938. Die Vogelbilder zur Historia Naturalis Brasiliae des George 
Marcgrave. 7. Orn. 86: 74-106. 


Books Received 261 Bull. B.O.C. 1998 118(4) 


Sibley, C. G. & Monroe, B. L. 1990. Distribution and Taxonomy of birds of the World. 
Yale Univ. Press, New Haven. 

Teixeira, D. M. 1992. As fontes do paraiso—Um ensaio sobre a Ornitologia no Brasil 
Holandés (1624-1654). Rev. Nord. Biol. 7: 1-149. 


Address: José Fernando Pacheco, Departamento de Zoologia, Istituto de Biologia, 
UFRJ—21944-970, Rio de Janeiro, RJ, Brazil. Bret. M. Whitney, Museum of 
Natural Science, 119 Foster Hall, Louisiana State University, Baton Rouge, 
Louisiana 70803, U.S.A. 


© British Ornithologists’ Club 1998 


BOOKS RECEIVED 


del Hoyo, J., Elliot, A. & Sargatal, J. (eds) 1991. Handbook of the Birds of the World. 
Vol. 4, Sandgrouse to Cuckoos. Pp. 679, 70 colour plates, c. 250 colour photographs, c. 
850 distribution maps, c. 8000 bibliographical references. Lynx Edicions, Barcelona. 
ISBN 84-87334-22-9. £110. 310 x 240 mm. 


This volume is the fourth in the beautifully illustrated HBW series (now one third 
complete). It begins with a lengthy Foreword by Jurgen Hafter, discussing species 
concepts and species limits in ornithology, dealt with under the general headings of 
typological species concept, biological species concept and evolutionary and phylogenetic 
species concepts. Examples are given illustrating clearly the differences in interpretation 
under the different species concepts, followed by a brief review of the increase in number 
of bird species described during the past 250 years. 

This Volume covers the four orders Pterocliformes, Columbiformes, Psittaciformes 
and Cuculiformes. The cockatoos are separated in a family of their own (Cacatuidae), 
rather than being lumped with the parrots (Psittacidae), so that six families are included: 
Sandgrouse, Pigeons and Doves, Cockatoos, Parrots, Turacos, Cuckoos. The resulting 
837 species described are more than in any of the other non passerine volumes. Awkward 
nomenclatural problems are described with particular reference to the tiny Fig-parrots. 

The considerably increased number of figures included in the 70 colour plates of the 
present Volume (compared to earlier volumes) has resulted in a significant increase in the 
number of contributing artists, with their associated differing styles. The editors hope, 
however, that the number of artists will stabilise for future volumes but are conscious of 
the desire to ensure that HBW continues to appear on schedule. 

In the first three volumes of the series the distribution maps (in conjunction with the 
text) are very useful and indicative but occasionally somewhat difficult to interpret, even 
for those regions with which I am familiar. The maps in Vol. 4 follow the same criteria, 
but the editors admit that the maps continue to present problems, available distribution 
data being frequently rather sparse and occasionally contradictory. Another difficulty is 
the adequate depiction of migration tendency, the solution adopted generally being to 
map the entire distribution area as green (signifying both breeding and non-breeding 
areas), with explanations being given in the Movements section of the text. Altitudinal 
movement is essentially impossible to indicate because of the small scale. In a few cases, 
areas are indicated for introduced species. 

As in previous volumes, the accepted French, German and Spanish names of species 
continue to be given, recommended Spanish names continuing to be published 
periodically elsewhere. The editors acknowledge the many museums and libraries which 
continue to contribute greatly to the healthy progress of the project. The very extensive 
bibliography contains many up-to-date references, just to quote one example—Tauraco 
ruspoli/Ruspoli’s Turaco includes comments in the Status and Conservation section from 
Borghesio 1997 (Bull. Brit. Orn. Cl. 117(1): 11-16) as well as being categorised as 
Endangered: Cites II. 

Pre-publication literature associated with Vol. 4 indicates that by means of a series of 
supplements (the first of which is in preparation) the up-to-date information on all 
species presented so far will be renewed indefinitely, including colour plates of new 
species described since publication of the corresponding volume. I find this extremely 


Books Received 262 Bull. B.O.C. 1998 118(4) 


encouraging, as has been the whole project to date and the editors (as well as BirdLife 
International) are to be warmly congratulated on the timely progress of this magnificent 
series. 

Lynx Edicions titles may be purchased from specialised bookshops or directly from 
the publishers: Lynx Edicions, Passeig de Gracia 12, 08007 Barcelona, Spain. Tel: 
+34 93 301 07 77; Fax: +34 93 302 14 75; E-mail: lynx@hbw,com; INTERNET: 
http://www.hbw.com 

S. J. Farnsworth 


Henry, G. M. 1998. A guide to the birds of Sri Lanka. Third edition, revised by 
T. W. Hoffmann, D. Warakagoda & U. Ekanayake. Pp. xlvit+488, 29 colour and 3 


monochrome plates, many line drawings. Oxford University Press. ISBN 0 19 563813 1. 
fS 22 SONS em. 


This third edition of Henry’s guide (the first in 1955, second in 1971) has been revised 
and updated by members of the Ceylon Bird Club, involving the addition of species, two 
new plates (by Bruce Henry, the original author’ s elder son) and a revision of the 
taxonomic treatment of species according to more modern texts. This involved the 
re-writing of the introductory sections to some Orders. Most of the species accounts 
remain as in the earlier editions, as do Henry’s original plates and line drawings, although 
the plate captions now give the page on which the species is described. Where necessary, 
species accounts have been revised, but the editors emphasise that for many species their 
distribution remains as Henry first described. Where species have been added to the Sri 
Lanka list, descriptions and details of occurrence are given but many are not illustrated. 
This guide thus retains the character of its antecedents and differs markedly from more 
modern field guides with their more terse styles. Nevertheless, this book provides more 
information on the birds’ natural history than many guides with which it will have to 
compete, and this may preserve for it a niche in the market place. 

Chris Feare 


Stattersfield, A. J., Crosby, M. J., Long, A. J. & Wege, D. C. 1998. Endemic bird areas 
of the world. Priorities for biodiversity conservation. Pp. 846, numerous maps, tables 
and photographs, some in colour. BirdLife Conservation series No. 7, BirdLife 
International, Cambridge. ISBN 0 946888 33 7. £37, $60. 24.5 x 17 cm. 1.7 kg! 


This book represents a milestone in many respects in the ornithological and 
conservation literature, reporting the findings of a major BirdLife project involving a 
huge number of people from all parts of the world. 

Over a quarter of the world’s bird species occupy small areas of the globe and this 
confers on these species a degree of vulnerability. In 1987, BirdLife (then ICBP) 
embarked on a project to catalogue and map the distribution of “restricted range” 
species, to record overlap with other bird species in this category and thereby to identify 
hotspots of occurrence of such species. The logic of this approach was that the limited 
resources available for conservation action could be targeted at a disproportionately high 
proportion of potentially vulnerable species in highly localised geographical areas. 
During the course of data collection for the identification of what are now termed 
Endemic Bird Areas (EBAs), studies of non-avian taxa revealed a degree of overlap 
between the distributions of their restricted range species and those of restricted range 
birds. Thus at least some EBAs have wider significance for biodiversity conservation. 

Eleven years after the initiation of the project, we now have this magnificent book 
which documents the 218 EBAs that were defined on the basis of having at least two bird 
species entirely confined to the area. Each EBA is allocated a number and an introductory 
box shows location, mentions key habitats and threats and tabulates the number of 
restricted range species that occur there. General accounts of the EBA, its restricted 
range species and threats are accompanied by tables showing the distribution, habitats 
and conservation status of the species, and by monochrome photographs (of birds, 
habitats and other animals) and plates of some of the birds. In addition to the EBAs, 
‘“‘secondary areas’’ are identified and described briefly in a separate section; secondary 
areas do not qualify as EBAs as fewer than two species are entirely confined there. 

In addition to these accounts, seven introductory chapters describe the background to 
the EBA concept, methodologies employed, a variety of data analyses illustrating how 


Books Received 263 Bull. B.O.C. 1998 118(4) 


these may be used to set priorities for targeting species and areas, and assessing 
conservation needs. The final chapter introduces the six regions under which the EBAs 
are treated, illustrated by maps, tables and colour photographs. 

All of these chapters have boxes which summarise key information. This, together with 
a two-page summary which presents the “‘meat”’ of the book with eye-catching headings 
and bullet-pointed facts, helps to make the book accessible to a wide readership— 
the promotional literature claims that the book is essential for conservationists, policy 
makers, scientific advisors, ornithologists and birders—but this approach renders the basic 
information accessible also to the popular media and, most important in many EBAs, 
schools. 

On might be tempted to regard this book as the pinnacle of BirdLife’s achievement in 
relation to EBAs but, as the authors point out, biodiversity conservation must change as 
animal and plant distribution change, and especially as our interpretation of taxonomy 
changes. In relation to birds, re-analysis of species concepts is leading to the elevation of 
many races to specific status and this will lead to the designation of further EBAs, 
especially through the elevation of “‘secondary areas’ to EBA rank. BirdLife, and 
especially the authors, are to be congratulated on the fulfilment of the initial stages of this 
major biodiversity conservation project and the publication of this excellent, accessible 
book which will have wide application. We should not forget, however, that over 70% of 
the world’s bird species are not confined to EBAs, and that even some of the most 
widespread species are undergoing catastrophic declines. Concentration on the EBA 
approach should not allow these species to be neglected, since they may well be indicative 
of more widespread depletion of biodiversity. 

Chris Feare 


Gaston, A. J. & Jones, I. L. 1998. The Auks. Pp. xviiit349, 8 colour plates, numerous 
text-drawings and figures. Oxford University Press ISBN 0-19-854032-9. £40, 
25.5 < 20 cm. 


The Auks are one of the most extraordinary and diverse of all bird families, and with 
the group being of a manageable size—only 22 extant species (or 23, see p. 191)—they are 
ideal candidates for a volume in the series Bird Families of the World. Auks may live in 
colonies numbering millions, can match the level flight speed of a peregrine falcon, can 
dive to 200m and catch their prey in virtual darkness, and may exhibit extraordinary 
ornamentation. The fascination and mystery of this group has obviously enthused the 
authors, who are responsible for a substantial portion of the research described, and 
whose authoritative writing reveal their comprehensive knowledge of auks. ‘The text is 
complemented by Ian Lewington’s superb colour plates which capture the nature of auks 
better than any I have seen previously. The result is a book which makes an outstanding 
contribution to an impressive series. 

The format is similar to that of previous volumes. The first half of the book comprises 
chapters on general auk biology, including systematics and evolution, distribution and 
biogeography, the role of auks in ecosystems, social behaviour, chick development, 
population biology and conservation. These chapters aim at synthesis, with an emphasis 
on trying to understand the remarkable diversity of life history strategies and ecology 
found among auks. The second part of the book comprises species accounts for the 
22 species of extant auks and also for the most recently extinct member of the family, 
the Great Auk. Each account includes sections on description, range and status, habitat 
and feeding ecology, displays and breeding behaviour, life cycle and population 
dynamics. The text is highly readable and includes numerous figures, tables, sketches 
and photographs. The text is also extensively referenced and throughout the book the 
authors find the correct balance between proven facts and informed speculation. A 
book of this nature also serves to highlight the fact that although tremendous advances 
have been made in recent years in many aspects of auk biology, e.g. our knowledge of 
many of the Pacific auks and the role of auks in the marine environment, there are also 
many areas of ignorance, including the basic biology of species such as Craveri’s 
Murrelet. 

This book is essential reading for anyone with more than a passing interest in seabirds, 
and one that I would recommend highly to any ornithologist. 

Ben Hatchwell 


Books Received 264 Bull. B.O.C. 1998 118(4) 


Ash, J. S. & Miskell, J. E. 1998. Birds of Somalia. Pp. 336, 5 colour plates, 3 colour 
maps, numerous monochrome photographs and maps. Pica Press, Mountfield. ISBN 
1-873403-58-5. £40. 253 x 190 mm. 


Extending from the East African equator to about 12°N, and with the longest coastline 
in Africa, Somalia is a country of immense ornithological interest. It embraces a large 
part of the unique Somali-Masai biome and occupies the eastern flank of a major 
migration flyway. Yet the documentation of its birds has until now been patchy. Early 
information from the northwest, the former British Somaliland, was brought together in 
the four volume work of Archer and Godman (1937, 1961), while pioneering explorations 
in the south were described mainly in Italian publications, including the multi-volume 
treatise by Moltoni and Gnecchi Rusconi (1940-1944), unfortunately interrupted by 
World War II. It was not until 1983 that John Ash and John Miskell produced, as a 
supplement to Scopus, the first annotated checklist to cover the whole of modern Somalia. 
These authors, resident at Mogadishu between 1978 to 1985, visited many areas, 
including the far north and northeast, and added over 50 species, including a new lark, to 
Somalia’s list. They began work on a national distributional survey, and this has resulted 
in the present book, an atlas which updates and greatly expands the information given in 
the 1983 list. 

An extended introductory section includes an historical review, and chapters on 
vegetation and various physical aspects of Somalia, with contributions from several 
specialist authors. The occurrence of migrant birds, Palearctic and Afrotropical, is also 
reviewed, and breeding seasonality discussed. These topics tend unfortunately to be dealt 
with in isolation, and more discussion of the impacts of vegetation and climate on bird 
distribution, movements and seasonality would have been welcome. 

The main body of the book deals with the 654 species recorded on the main list, which 
includes 151 Palearctic migrants and 7 endemics. Nomenclature and sequence broadly 
follow The Birds of Africa. Excellent quality maps plot occurrence and known breeding 
distribution on a 1/2° x 1/2° grid. The accompanying text gives details of status, 
abundance and habitat, all Somalia races being mentioned. Breeding information 
includes seasonality and clutch size. For scarce or little known species all records are 
given. Further lists detail offshore records, hybrids, and species requiring further 
confirmation. An extensive gazeteer provides modern Somali as well as English or Italian 
names, and should prove very useful to ornithologists and others. A full bibliography 
includes over 500 citations. Five lively coloured plates by Martin Woodcock portray races 
and species special to the region. 

This is an attractive and well-produced book, essential for anyone with an interest in 
Somalia’s wildlife, or indeed with a wider interest in East African bird distribution or 
migration. It provides a thorough review of current knowledge of Somalia’s bird 
populations and distribution, at the same time indicating areas where much still remains 
to be discovered. It will certainly provide the basis and stimulus for further exploration of 
the country when access improves and more settled conditions return. 

David Pearson 


NOTICE TO CONTRIBUTORS 

Papers are invited from Club Members or non-members, especially on taxonomic 
and distributional topics; descriptions of new species are especially welcome and 
may be accompanied by colour photographs. Two copies of manuscripts, typed on 
one side of the paper, double spaced and with wide margins, should be sent to 
the Editor, Prof. Chris Feare, 2 North View Cottages, Grayswood Common, 
Haslemere, Surrey GU27 2DN, UK. All contributions, including In Brief 
articles, should follow the style of main papers in this issue of the Bulletin. 


A contributor is entitled to 10 free offprints (16 if 2 or more authors) of the pages of the 
Bulletin in which his/her contribution, if one page or more in length, appears. Additional 
offprints or offprints of contributions of less than one page may be ordered when the 
manuscript is submitted and will be charged for. Authors may be charged for proof corrections 
for which they are responsible. 


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annual subscription, should be made to the Hon. Secretary (address as below). 

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NUMBERS OR OTHER PUBLICATIONS 

The Bulletin (for 1999 onwards), together with annual index, may be purchased 
(postage free) by Non-member Subscibers on payment of an annual subscription of 
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similarly be obtained, on request to him. 


PAYMENTS 

All amounts quoted are net and should be paid in £ sterling, if possible. Payments 
in other currencies must include a further £4 for UK bank charges (except for 
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Eyebrook, Oldfield Road, Bickley, Bromley, Kent BR1 2LF. 


CORRESPONDENCE 
Correspondence on membership, changes of address and all other matters should 
be addressed to the Hon. Secretary, Cdr M. B. Casement, OBE, RN, Dene Cottage, 
West Harting, Petersfield, Hants GU31 5PA, U.K. For details of Club Meetings see 


inside front cover. 


Registered Charity No. 279583 


CONTENTS 


CEUB NOTICES foo. oes eee el al, sree reeurtay ai etenear acerca ye Peiranleyioet tary) teblietst cits al alta ate ana er 


LAMBERT, F. R. & RASMUSSEN, P. C. 


A new Scops Owl from Sangihe Island, 


Indonesia’. 5 60. Soe ee ee ae eh ce) ol ae 


LECROY, M. & PECKOVER, W. S. 
HOLMGREN, J. 


WELLER, A.-A. 


I. Amazilia Sumichrasti Salvin, 
biogeography within the A. beryllina complex 


OLSON, S. L. 


Misima’s missing birds 


ee 68 we 6 et) ® 6S ee lee ee 


A parsimonious phylogenetic tree for the swifts, Apodi, 
compared with DNA-analysis phylogenies 


o@ © © 8 e 8 6 6) a eee) wee 


On types of trochilids in the Natural History Museum, Tring. 
in relation to morphology and 


e 8 6) © es ee fe pis) ea ee oe 


Lectotypification of Charadrius rubricollis Gmelin, 1789 ..... 


PACHECO, J. F. & WHITNEY, B. M. Correction of the specific name of 
Long-trained: Nightjar s; 2. De kk ee 
BOOK ‘RECEIVED’. 6) 6505522) ot oe ok Be am 1 ae ha de a 00) 


259 
261 


The Bulletin is despatched from the printers on publication and is sent by Surface Saver 
Postal Services to all European destinations outside the U.K. and by Air Saver Postal 
Services to destinations outside Europe. Those whose subscriptions have not been received 
by the beginning of a month of publication will have their copies despatched by surface 
mail, after their current subscription has been paid. 


COMMITTEE 


Revd T. W. Gladwin (Chairman) (1997) 
Mrs A. M. Moore (Vice-Chairman) (1997) 


Cdr M. B. Casement, OBE, RN 
(Hon. Secretary) (1996) 


D. J. Montier (Hon. Treasurer) (1997) 


Hon. Editor: Prof C. J. Feare 


D. Griffin (1997) 
N. J. Redman (1997) 
Mr R. E. Scott (1998) 


Chairman of Publications Sub-committee: Dr R. P. Prys-Jones 
Publications Officer: S. J. Farnsworth 


Printed on acid-free paper. 


Published by the BRITISH ORNITHOLOGISTS’ CLUB and printed by 
Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 


R. P. Prys-Jones (1995) 
N. H. F. Stone (1995) 

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